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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
writing from the author.
ISBN: 978-1-4349-9074-7
Printed in the United States of America
First Printing
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Or visit our website and online catalogue at www.rosedogbookstore.com

ACKNOWLEGMENTS
I want to express my love and gratitude to my wife and my daughter for all the remarkable effort
designing the pictures and illustrations; without their help this book would not be possible. My
son, whose encouragement and support is invaluable. To the loving memory of my father; he
taught me so many things about birds.
Special thanks to all those canary fanciers that I have met during times of my life, and from
whom I have learned so much. I am especially indebted to Antonio Sanchez Bermudez, friend
and colleague. He passed away long ago; a great lost for all those who learned so much from his
wise advises. Alfredo Rovere generously provided valuable information in the 70s, when specialized canary magazines were not affordable for me. Dr. Lidia Bermudez has been a tremendous support in the completion of this book. Silvio del Valle and Maria Elena Rodriguez were
the pioneers in the first attempts to get canary pictures for my book. Karelia Llanes was of a
great help in computer assistance.
My former teachers in all courses of Genetics and Animal Genetics at the University of Havana,
Faculty of Biology, implanted in me the unbreakable decision to go deeply in knowledge, no
matter how hard it might seem to be at first.

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CONTENTS
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . iii
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii
Chapter 1: Basic Concepts on Genetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Chapter 2: Mating Systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Chapter 3: Pigments and Feathers in the Canary. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Chapter 4: The Dominant White and the Recessive White Canary . . . . . . . . . . . . . . . . . . . . . 17
Chapter 5: The Yellow and the Lemon-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 lipochromes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 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 Simple Pairing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
Chapter 16: Concomitant Factors in Color Canaries: other mutations . . . . . . . . . . . . . . . . . . 76
Chapter 17: The Nomenclature of Color Canaries . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
Chapter 18: Quantitative Canary Breeding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83

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 15th or
16th century. At that time, that melodious little bird created a very special predilection among
European aristocrats. The canary rapidly spread all over Europe, and consequently variations
soon appear in its genome. It seems that at the end of the 17th century two important mutations
took place, dominant white and yellow. Surprising avalanches of mutations 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 knowledge. Is each and every canary breeder in such position and possibility? Im 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 many articles available on the internet. What I have found as my own conclusion is that, no possible flawless application could be
achieved if the reader is not prepared in basic genetics, animal breeding techniques, and a proper control of what he/she is doing during the breeding season, no matter what the complexity
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 amount of
money 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 worthwhile? Furthermore: Is it the merit of the buyer obtaining good and excellent birds next season,
or is it the merit of the seller who bred the parental strain? The answer is simple: the seller is.
In my own opinion, it is better for any fancier to apply as much knowledge as possible, since this
is the core of canary breeding. Better birds based on better knowledge. If you have only regular birds, and you apply technical knowledge properly, it is possible to obtain outstanding birds
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in two or three breeding seasons. Youll be proud to say: those are my very own birds, I made
them myself. You will be satisfied without spending a lot of money, and those birds being better 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 more) to accomplish canary breeding in a modern, technical, and methodological
way. There are not special secrets in obtaining excellent birds. Applying knowledge properly is
the answer. Canary breeding is only a particular specialty in animal breeding. High performance results are based on this. Hence, specific procedures might be taken into account.
Some simple suggestions are mandatory to those readers that take this book having in mind to
thoroughly learn and apply its content. First of all, it is a must to understand the content of the
first chapter, sine qua non, it is impossible to go further. Besides, it is important that every single chapter be analyzed carefully, in order to master its content in full. I do not advise reading
a chapter if the previous one has not been thoroughly understood. These remarks cannot represent a discouragement at all; on the contrary, smart precautions that guarantee the proper
development of your knowledge.
The author

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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 material is included. Only basic structures of the cell will be considered, not beyond the concerns of basic Genetics.
Every single tissue, in any animal or plant, is built by microscopic structural and functional
units called cells. They show different sizes and shapes, depending on the function accomplished. As an example we can mention the kidney cells which are completely different from the
brain cells, as they perform a very different function. In general terms we may say that a cell is
basically formed by a nucleus, nucleus membrane, cytoplasm, and cell membrane, as it is shown
on Fig. 1.

Fig. 1 Basic parts of an animal cell.


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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 filament structures, like microscopic
threads in a complex hank. The size, number and shape of chromosomes are different in every
animal species. The number of chromosomes in the cell nucleus is constant in every species. In
the chromosomes genes are located, which are the functional units bearers of the genetic traits
in all individuals. Between 1910 and 1928 well conducted investigations documented the linearity of genes in the chromosomes; it is, genes are placed in chromosomes as beads in a necklace: one after the other.

Cell Division
The production of somatic cells (cells of the body; from soma: body) is accomplished by a
process of cell division called mitosis (from the Greek mitos: thread). Each single cell is divided 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 mitosis different phases are considered: prophase, metaphase, anaphase,
and telophase; each one corresponding to important stages in cell division. Next, those steps
will be roughly explained.

Fig. 2 Two cell divisions in a regular mitosis process.


Prophase: is the starting point of cell division. Each single chromosome is longitudinally divided in two perfect halves, named chromatids; in fact they are strands resulting from chromosomal duplication. These strands are linked together in just one point called centromere, located
at one end of the chromosome, in the middle or in any other position between the center and
the extreme end of the chromosome, as shown in Fig. 3a. Toward the end of the prophase the
nucleus membrane disappears, and chromosome are grouped in the equatorial plate of the cell.
The so called mitotic spindle is formed. This is the beginning of the next stage, the metaphase,
as shown in Fig. 3b.

Fundamentals of Color Genetics in Canaries

Fig. 3 Mitosis process in an animal cell.


Metaphase: the mitotic spindle is completed, and chromosomes are placed just in the middle of
that structure coincident with the equatorial plate of the cell. Now, each chromosome is seen as
a pair of chromatids, joined by the centromere. At the end of the metaphase the centromere is
gone, but chromatids are still one next to the other.
Anaphase: now the chromatids set apart and migrate to each pole of the mitotic spindle. In fact,
they are true and individual chromosomes, as shown in Fig. 3c. From now on they are genuine
chromosomes. We may realize that a fundamental part of cell division is the own division and
duplication of chromosomes, themselves.
Telophase: the mitotic spindle disappears, and chromosomes become much 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 complete division, and creating two daughter cells
which bear the same amount of chromosomes as the mother cell, due to the migration of chromatids-in fact, true chromosomes at this time- to opposite poles of the mitotic spindle being
included in equal amounts in both daughter cells. See Fig. 3e.
All of the above, as was mentioned before, is regarding somatic cell division. Something different occurs when division involves the creation of gametes egg-cells and spermatozoids- which
is known as meiosis or reduction division. Germ line cells only bear half of the amount of chromosomes compare with somatic cells. If the gametes of a certain animal species bear n chromosomes, then its somatic cells bear 2n chromosomes, known in Biology as the diploid number, while the gametes own a haploid number n. The reason for haploidy and diploidy phenomena shall be understood at the end of the following.

Octavio Perez-Beato
Meiosis
During the normal reproductive animal cycle, testis and ovaries play an outstanding role forming 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 completed, 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 particular 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 chromosomes, which in turn will create two cells, being each one
loaded with only one single group of chromosomes. Nevertheless, we may observe that these
chromosomes are longitudinally differentiated in two chromatids. Fig. 5 shows the first and second meiotic division. In the last one each chromatid is separated from the other, and in turn
becomes an individual true chromosome, 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 spermatozoids, accordingly.

Fig. 4 In (a) chromosomes are one next to the other. In (b) there is one single chromosome of
each pair in each group; typical meiotic metaphase.
All this biological process has been explained in a very simple and schematic way on behave of
simplicity and easy understanding, but the biological reality is so much more complex.
As you may see in Fig. 5, not four egg-cells are produced at the end of the second meiotic division, only one egg, instead. The other three cells undergo a degenerative process and stop
development. On the other hand, it is not the case regarding the four spermatozoids since they
really complete full development, from each cell that reaches meiotic metaphase. It is also
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Fundamentals of Color Genetics in Canaries


important to take into account that in meiosis two cell divisions are accomplished, but only
one single chromosome division is performed.
Now, one single question arises: What is the objective of the meiosis process to allow spermatozoid 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 biological being will bear in each and single somatic cell the same amount of chromosomes as its parents, 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 male and female gametes.


The cell formed with these contributions is called zygote, and of course will bear the same
amount of chromosomes as is customary in any individual of its same species. Such a process is
clearly shown in Fig. 6.

Fig. 6 The approach of a spermatozoid to an egg-cell is depicted, together with the inclusion of
the spermatozoid inside the egg mass.
5

Octavio Perez-Beato
The contribution in terms of chromosomes from both gametes readily creates the zygote. Soon,
the zygote will start mitotic division, and a new living thing is formed here on. This new individual has inherited traits from its parents, loaded in the chromosomes located in each and
every one of its cells, which are nothing but replications of the original chromosomes from the
spermatozoid and egg-cell, from which the zygote was originated.

Terminology and Definition of Some Basic Genetic Concepts


As it was demonstrated in previous paragraphs, chromosomes are paired in every somatic cell,
and in an unpaired fashion in the gametes. If we recognize the existence of pair of chromosomes, this means that each pair is carrying a whole series of genes, that in turn are also located by pairs on specific locations along the chromosome, except in the chromosomes called XY.
From this, it is quite simple to assume that there are specific locations along each single chromosome where genes are placed. The specific location for a gene on any chromosome is known
as locus; plural loci. Each single gene located in the same chromosome pair, is called allele.
In modern terms and simple way, the gene must be defined as a factor that determines 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 following: 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.

Fundamentals of Color Genetics in Canaries


Dominance, Expressivity, and Penetrance of a Trait. Phenotype and Genotype.
It is said that an allele is dominant regarding the other allele in the gene, when the heterozygote
for example, Bb- only shows the trait determined by the allele B.
Then we may say that b is recessive regarding B. Consequently, all individuals bearing BB or Bb
are no different at a glance. Nevertheless, BB is homozygous regarding trait B, and Bb is heterozygous, since b is the other allele in this pair. Taking this into account we simply reach the
concept of phenotype, which is precisely what we observe externally in any living thing. This
mean that based on phenotype only, BB and Bb are identical, since both show the same trait B
(dominant trait), but from a genetic point of view they are different, since Bb bears the b allele,
and BB does not. Thats why we might say that their genotypes genetic types-are different. Not
always dominance is shown in this simple 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 may
say that it is an intermediate dominance. Some other times the dominance is incomplete, since
the recessive allele gets some expression on the phenotype. Furthermore, we have to recall two
important phenomena regarding any specific trait. It is possible that a dominant allele will not
be expressed the same all over the population. Some few differences are readily seen from one
bird to the other. These ranges of signs are called variable expressivity. Such is the case regarding the frost character in canaries. We may see birds with a mild frost up to heavily frost. This
range of appearance is telling about a variable expressivity.
The other phenomena that I want to mention is the reduced penetrance which refers to the proportion of birds expressing a particular trait. For example, let us suppose we have 50 canaries
bearing a certain genotype; only 40 show the corresponding phenotype. Consequently, the penetrance of the trait is 40/50 = 0.80; this means penetrance = 80 %.
We ought to consider that expressivity and penetrance are both affected by environmental factors such as: temperature, 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 chromosome X, we say that the inheritance of the
trait is sex linked. That is why a male canary could be heterozygous or homozygous for any sex
linked trait, since the genetic formula for a male canary is XX and then the male may carry the
same allele in both chromosomes, or different ones in each X chromosome for any particular
trait. On the other hand, the genetic formula for the female canary is XY. On the Y chromosome
there is not any allele of our concern, and we usually say that Y chromosome is empty. Then
again, for a female it is only possible to bear sex linked traits (genes) on the only one X chromosome that she possesses. It is clearly seen that a female canary never ever could be heterozygous for any sex linked trait.
The opposite situation is for the traits located in non sex chromosome and then we say the specific trait autosomic.

Octavio Perez-Beato
Proportion of Genotypes Expected in a Particular Mating
It is of the most importance for any fancier to precisely predict the genotypes of descendants,
considering the genotypes of the selected parents. Obviously, fanciers in general are only interested on one, two or three traits in the breeding accomplished. Hence, all efforts are oriented
on the calculation to accurately predict the descendants genotypes. Let us assume that a mating is accomplished between an AA genotype male canary and an Aa genotype female 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 accomplished by making a simple chart
widely used in genetics, known as the Punnett squares.

Fig. 8 Example 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 columns are depicting the female canary gametes one allele per
gamete-and then rows represent the male canary gametes. Each cell represents the allele input,
one from the male and one from the female. 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 means that 50% of the offspring will bear genotype AA. This is a simple hypothetic example 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: male canarys genotype Aa
BB; female canarys 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 former example, and as in any other case, each single cell represents a particular genotype which could be obtained in the expected offspring.

Fundamentals of Color Genetics in Canaries


Based on this, if we are interested in obtaining the male canary genotype in the offspring for
both involved traits, the information from the Punnet squares will provide us with the expected proportion of that specific genotype in the expected offspring. As we got two cells with genotype 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 same genotype as the male
canary regarding the considered alleles. In the upcoming chapters the reader will learn the alleles comprising the genes involved in the color inheritance in the canary.
Using the Punnet squares the fancier can make 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 make clear some few aspects, for those readers
that are not familiar with genetics; it is, if the total expected genotypes in the offspring is 4, 5 or
more, and the number of chicks born is small, chances are that some genotypes will not appear
in the expected offspring, since all these proportions are based on probabilities.

CHAPTER 2
Mating Systems
There are two basic mating systems: inbreeding system, in which related birds are bred, and out
breeding system in which not related birds are used in breeding. Besides, the relation might be
a close one, or a collateral one, depending on the purpose of the fancier, and the breeding possibilities based on the available stock. Below, the scheme for such mating is offered as a guide
for the fancier to accommodate the objectives of breeding procedures. First, it is important to
establish some simple definitions to better understand what is printed in the scheme below.
Line-breeding: mating in the
direct line of descent. In linebreeding, what is usually
obtained is a discrete homozygosis for selected traits.
Lineage: group of individuals belonging to a specific
breed (in our context: color
anaries) which are homozygous for certain traits.
Breed: a specific group of
individuals which bear a relative homozygosis for specific traits, but that homozygosis condition is more discrete than in the lineage.
Sometimes, it is difficult to
establish a definition between
lineal or lineage condition,
since lineal relation is a genet10

Fundamentals of Color Genetics in Canaries


ic trend toward lineage. Inbreeding tends to homozygosis. Then, if we are interested in creating
a lineage we ought to use inbreeding procedures.
Since inbreeding creates homozygosis, it is expected that recessive traits not frequently seen in
the population will suddenly appear, just because now they are in a homozygous 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
mandatory. We cannot be tolerant, not with the slightest fault. The most meticulous 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, some cases of sterility may suddenly appear, and other disturbances related to reproductive traits. Fortunately, experiments have demonstrated that, normal
reproductive levels are recovered as soon as we start mating inbred lineages among themselves.
On the other hand, out-breeding leads to heterozygosis which in turn produces a phenomenon
known as heterosis; it is the counterpart of depression due to inbreeding. We may say that heterosis is present when mating 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 outcome of a bird sharply superior
compare with its parents. This is heterosis.
Breeding not related individuals has the disadvantage of possible introduction of traits not convenient to our goals. Furthermore, it is highly dangerous from a genetic point of view to introduce new birds, if the fancier is already working to establish a lineage for certain specific traits.
Next, some mating schemes are shown for the breeder to have an illustrated reference, in order
to set up breeding programs according to what is available and possible.
Fig. 10 shows mating that start with male (1) and female (2), producing an offspring (3), which
bears 50% from the father (1), and 50% from the mother (2). All males from group (3) will be
selected according to what the breeder wants; the best of those males will be back-crossed to
female (2), then obtaining offspring group (4), which possesses 75% from her mother (2); the
best male from group (4) will be back-crossed again to female (2) obtaining the offspring group
(6), which possesses 87.5% from female (2).
Then again, the best male from group (6) will be back-crossed to female (2) to create the offspring group (8), which now possesses 93.75% of the phenotype from female (2). This group (8)
is almost a copy of female (2), very near to a clone. Using this mating procedure we replicated
the phenotype of female (2), in approximately 94% in a whole group of descendants.
What is said before regarding female (2), is true for male (1) represented in the left hand side of
the scheme and fully developed in a parallel fashion. In this way the breeder will have descendants in both lines of the breeding lineage: maternal and paternal, which is mandatory for the
continuity of the breeding system and genetic improvement, as well.
11

Octavio Perez-Beato
Finally, the best birds from group (8) will mate the best from group (9), to obtain group (10) offspring, which is similar to the first offspring (group 3) from male (1), and female (2). Besides, if
group (10) mates to group (8) and (9), then we will obtain groups (11) and (12), which turn to
be quite similar to groups (4) and (5) from previous years, since group (4) and (5) represent
24/32 of female (2) and male (1) genotype, respectively, and then groups (12) and (11) represent
23/32 from female (2), and 9/32 from male(1), respectively.
We may continue mating groups in analogous form, taking into account that birds from group
(8) are very similar to female (2), and birds from group (9) are very similar to male (1). Keep on
breeding in a similar way to obtain groups similar 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 similar considerations.
It is of the most importance to start with birds carefully selected, and if possible of known pedigree, no matter what the linebreeding scheme is applied, remember that inbreeding itself is a
tool to surface faults, and then again culling might go to great length if the mating starting-line
is not attempted with good quality birds.

Fig.10 Inbreeding scheme to obtain


nine offspring groups

Fig. 11 Inbreeding system where


males A and B are siblings,
and females C and D are not related.

12

Fundamentals of Color Genetics in Canaries

Fig.12 Both original females are


not related. On the last mating a
heterosis effect must be evident
in the offspring. Arrow heads are
used as in the previous figure.

13

CHAPTER 3
Pigments and Feathers in the Canary
The plumage color in the canary is biologically conditioned by means of two pigments: the
melanins and the lipochromes. The combination of these two substances in the feathers of the
canary, give birth to all combinations that we admire today in our beloved feathery pet. Besides,
melanins and lipochromes are genetically affected by other factors, which will be the topic of
upcoming chapters. These substances are fully studied by the science of Biochemistry, which is
in charge of the research and study of all the chemical processes in living things.

The Melanins
The canary bird possesses two types of melanins, though some authors state that are more than
two. For the simplicity of explanations, and better understanding of basic concepts on this
topic, I will consider only two melanin types throughout the book. These melanins are: eumelanin or black melanin, readily seen in Green, Bronze, and Blue intensive canaries. The
pheomelanin (brown melanin) is a pigment very well seen in Yellow Brown and White Brown
canaries, among others. These melanins are biochemical elaborated by the canary itself, based
on the genetic information of the bird. It is not enough that a canary could elaborate its own
melanins, it is mandatory that very special and specific substances have to be present in this
biochemical process. These substances are known as oxidative enzymes oxidases-, which act
on the melanin, and then developing the appearance of each type of pigment. If this enzyme
would not be present to carry on the oxidative action, it will be impossible to see eumelanin and
pheomelanin in the plumage 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

Fundamentals of Color Genetics in Canaries


These pigments are present in the canary diet, then metabolized and fixed in its skin; as feathers are epidermal products any canary with the proper genetic information shall deposit
lipochromes on its feathers. Lipochromes range from yellow to red, and canaries get them by
means of the metabolic process of carotenes and xanthophylls, which are substances fully
embedded 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 many plants. The darker the green color the better as a canary food.
The genetic aspect regarding the way the canary transforms carotenes and xanthophylls into
pigments to color its plumage, will be considered in detail in the chapter about White Dominant
and Recessive White. As a previous example we may say that the Red Siskin has the necessary
genetic information to transform carotenes into red pigment, red lipochrome, which is deposited in the males feathers and on a discrete fashion, in the females, too. On the other hand, the
yellow canary is able to transform xanthophylls into yellow lipochrome, and then deposit it in
its feathers. I strongly agree with other authors that have stated four fundamental steps to set
up the proper color in the canary; these steps are:
a) Vegetables substances carotenes and xanthophylls- must be assimilated by the bird.
b) Those substances must be transformed into true pigments, due to the metabolic action
of the canary organism itself.
c) The pigments already elaborated must be concentrated in the feathers. In fact, these
pigments are concentrated in the skin of the bird; as I said before feathers are of epidermal origin; these pigments will be deposited in the primordial feather structures in
the skin. Finally, they are seen in the full-grown feathers, if the canary has the proper
genetic information to do so.
d) As the last step, other substances acting according to the genetic information present
in the genotype, will determine in what way and how much of those pigments will accumulate in the feather structures. The genetic aspect of these four steps will be explained
in the upcoming chapters.

The Frost Feather and the Intensive Feather in the Canary


There are two feather structures in the domestic canary; the frost feather and the intensive
feather. The frost feather presents the barb extremes with no lipochromic pigmentation at all,
which make the plumage to appear like frosted. At the same time this kind of feather is soft, like
a down. On the other hand, the feather known as intensive reveal lipochrome pigment up to
edge of the barbs, being shorter and more rigid than the frost feather.
The frost and intensive feathers are controlled by two alleles; the intensive is dominant, the
frost if recessive. This means that three different genotypes are possible regarding this kind of
feather structure. Let I be the dominant 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 mandatory to fully explain what to expect
15

Octavio Perez-Beato
in genotype II birds. This homozygous dominant is a bird with long fingers and nails, extremely 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 embryonic process; sudden death in the nest. If it survives, definitely will be a degenerated biotype. Besides all previous characteristics, its plumage 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 homozygous condition is sub-lethal. There is something else,
allele I shows an incomplete dominance, and variable expressivity since birds with genotype Ii
ranges from a true and harmonic intensive appearance up to birds with a very mild frost. These
might be excellent breeders but not recommended as show quality birds.
On the other hand, if two frost birds are mated, the offspring will be composed of excessively
feathered animals, rather lazy, and not really good in rearing their chicks. On the average, there
will be no problems regarding sub-lethal effects, but they do not match with any standard.

Fig. 13 Mating frost and intensive will yield an offspring comprising frosts and intensives
in the same proportion.
According to what was explained in the previous paragraphs regarding frost and intensive
feathers, mating 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 dominant white is the most common 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 lipochromic tint (yellow,
orange or red) is apparent in the primary wing feathers, and sometimes extended to the coverts.
It is believed that this canary appeared at the end of the XVII century. It was a mutation. These
birds are white color just because they are unable to deposit lipochromes in its feathers, which
is determined by a gene F that, in heterozygous condition (Ff) produces a dominant white phenotype; if it appears in homozygous condition FF, the bird bearing such combination will die
during the embryonic stage. That is why it is said that, the gene that determines the dominant
white is lethal in homozygous condition. As a result, any dominant white canary is heterozygous
for that specific trait. This is a dominant autosomic gene. Nevertheless, there are other loci
where important genes involved in different metabolic steps regarding lipochromes, are located. I will explain in detail the theory that supports this genetic event.
Locus K
The dominant allele K allows the normal assimilation of carotenoids. The recessive allele k does
not allow the normal assimilation of these substances. All birds with genotypes KK or Kk will
assimilate the carotenoids on a normal basis. Unfortunately, the genotype Kk does not get
enough carotenoids to be lately transformed into yellow pigment. This aspect will be discussed
in the next chapter.
Locus G
The dominant allele G allows the conversion of carotenoids into yellow pigments.
On the other hand, the recessive allele g does not allow such conversion. All canaries with genotype Gg will readily transform the carotenoids into yellow pigment, though they are bearing the
recessive allele g. This allele g is just a convenience, and the theoretical homozygous 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 German white is a potential mutation. 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 1908. 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 1908 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 almost disappeared. Taking into account all of the above, it is obvious that the mutation responsible for the
recessive white canary has occurred several times in the past. This means that it seems 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 plumage of the recessive white does not show any trace of lipochrome, as is the case in the
dominant white canary. Besides, the skin of the recessive white canary is bluish color.
These birds as being unable to assimilate the carotenoids, cannot either elaborate the vitamin
A, which is a normal biochemical process in the rest of the canaries, from the carotenoids in
their food. This is the reason why it is necessary to supply vitamin A in the diets of the recessive
white canaries.

Genetic Characteristics
Based on all that has been discussed regarding the dominant white and the recessive white
canaries, now we are able to discuss the genetic formulas that characterized these birds.
The dominant white canary does not deposit lipochrome pigments in its feather structures,
except on the areas already discussed in previous paragraphs. But certainly does assimilate the
carotenoids, and also transforms them into lipochrome pigments. Then, according to the function of the previously explained loci in this chapter, now we have that a dominant white canary
18

Fundamentals of Color Genetics in Canaries


has the genetic formula: KKGGFf; and a recessive white canary is formulated as kkGGff. As I
have previously expressed, the allele F is lethal under homozygous condition; enough reason to
explain that mating two dominant white birds the obtained offspring will show a 25% of embryonic mortality. See the chart below:

Fig. 14 The offspring group (1) is homozygous for allele F, which is detrimental to the
embryonic development. The offspring group (3) are yellow canaries, which genetic formulas
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 homozygous
regarding allele F. Next chart shows the results:

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


of mating a Dominant White male and female.
Cell (1) represents homozygous FF, which will not complete their embryonic development. At
the same time this group represents 25% of the offspring. Besides, cells (2) and (3) represent
50% of the offspring which are dominant white canaries. Finally, cell (4) represents yellow
color offspring.
It is very important for the reader to understand every single detail explained in this particular chapter, since it is the basis to really understand the genetic principles regarding the
inheritance of classic white canaries. Besides, if the reader takes hold of these fundamental
concepts the following chapters will be easy to understand, mainly when reading will reach
all the theory and statements on the Red Canary, which is the most attractive approach to
lipochrome genetics.
19

CHAPTER 5
The Yellow and the Lemon-Yellow Canary
The yellow canary appeared as a mutation 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 biochemical action of carotenoids in the canary
plumage. Besides, in chapter 4 the genetic formulas of the white canaries have been established.
Now again, the yellow canary as a counterpart of the white recessive canary, fully assimilate the
carotenoids, in particular those pigments that will be transformed into yellow lipochrome.
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 formula of the yellow canary (KKGGff) was established as the result of mating two dominant white
pair. See Fig. 15 as a reference. As the allele K is dominant over allele k , it is possible to say
from a genetic point of view- that a yellow canary may also bear, theoretically, the genetic formula KkGGff, which will not affect the normal assimilation of carotenoids. This heterozygous
canary regarding locus K will show a poor yellow lipochrome in its plumage, since the dominance of allele K is incomplete then producing a partial assimilation 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 mandatory to consider a special point about locus
K. If we take a closer look to the previous formula KkGGff though it is obviously characteristic
of a poor yellow color plumage, it is also indicating a white recessive split bird, since it is heterozygous Kk. At this point of the analysis I particularly do not think it is the genetic explanation for the poor yellow color in the plumage. Nonetheless, it could be the genetic formula that
bore the parents of the white recessive birds that appeared in different times in the past, which
pointed to the possibility that the white recessive trait is a recurrent mutation, as was stated in
Chapter 4.

20

Fundamentals of Color Genetics in Canaries


Genetic Characteristics
The lemon yellow canary is widely known among canary breeders. This bird does not possess
different lipochromes than those bears by the common yellow canary. The special appearance
of its lemon-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 lemon-yellow color. This feather
structure is genetically controlled. Let us call allele s the one that determines such special structure, and S the allele for the common feather structure.
The allele s is known in canary breeding as blue optic factor. Taking into account what is known
for recessive alleles, a lemon-yellow color canary has to carry ss in its genotype to be a visible
lemon-yellow color bird. In other words, that canary has to be homozygous ss to show such a
color as its own phenotype.
As the reader surely knows the genetic formula of the yellow canary from a previous chapter, let
us add the optic factor gene to that basic formula to obtain the lemon-yellow canary: KKGGffss.
It is important to know that the allele S that determines the common structure of the feather,
acts based on an intermediate dominance, hence the heterozygous canaries Ss show an intermediate phenotype between common yellow and lemon-yellow, known as mid-lemons. It seems
that this denomination was formerly acquainted in France by the French term demi-citron
(half-lemon).
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 may 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 many backyards during the fall and winter migration. When it gets wet, the iridescent blue color vanishes, and the bird plumage turns into a rather dark brown color. The characteristic 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
momentarily altered in its surface, due to the water, and then is unable to properly reflect such
a color. What we see at that very moment is the true pigment underneath the surface structure,
since the blue color in birds, no matter what the species is, does not exist as such, it is always
the effect of that special structure on the surface, which may produce from a very light to an iridescent metallic blue color.
It is an interesting remark that the dimorphism apparent between the male and female of the
Indigo Bunting is due to this optic blue factor only present in the male plumage. Unfortunately,
our canaries do not posses such an efficient feather structure to reflect the true blue color, and
by the moment time we only can admire that mild blue optic factor in the lemon-yellow and
other colors that will be discussed in upcoming chapters.
Now, the reader can add another allele to the general genetic formula of the common yellow
canary, since it has been established the normal allele that denotes absence of the blue optic factor; the complete genetic formula is KKGGffSS for the common yellow canary, and for the
lemon-yellow is KKGGffss
21

Octavio Perez-Beato
From the mating of a lemon-yellow bird to a non-lemon-yellow, the offspring will be as shown
on Fig 16.

Fig. 16 Mid-lemon proportion offspring -100%- from a yellow and lemon yellow mating.
As readily seen in the previous chart, the whole offspring will bear the optic factor, and then
they will be mid-lemon birds. If the conducted mating is mid-lemon and lemon yellow, the offspring will be : 50% mid-lemon , and 50% lemon 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 autosomic trait, according 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 attempts to create a red canary were made. I may say, and I think I
am certain that it was Dr. Duncker the first to be interested about genetics in the canary.
Nevertheless, by means of Dr. Duncker himself it was known that the first canary breeder that
carried experimental mating with the red siskin (Spinus cucullatus) and the female canary was
named Mr. Adams, but the methodology to get the hybrids was stated by Dr. Duncker.
I also have to mention B. Matern and F. M. Durham,that made important contributions regarding 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 time full information regarding all concerns about the red canary.
I think it is necessary to make a review, though it might be simple, about theories regarding the
red canary genetics. According to Dr. Duncker theory the yellow lipochrome was absent in the
red siskin. Then in the mating of a red siskin and a yellow female canary, the hybrid will get the
red factor from the red siskin, and the yellow color from the female canary, being this combination responsible for the copper color shown in F1. Next it follows a back-cross F1 male with
a yellow female canary; the offspring came to be: copper color, pied-oranges, pied-yellows, light
oranges, and light yellows. The males were fertile, but not the females. Strikingly, dissections
made by Dr. Duncker on these F1 females, demonstrated and absent of reproductive organs.
In his book Mr. Gill provides a very interesting information from his own experience about the
F1 cross to female canaries; the figures are as follow:
Total mates
Total eggs laid
Infertile eggs
Dead in the shell
Total chicks hatched
Total chicks reared

52
215
171
4
40
30 (22 males, 8 females)
23

Octavio Perez-Beato
From this information 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 attempt to
hybridize two different species, in which the reproductive results are usually very low.
On the other hand, we may observe that the number of eggs per nest is on the average expected for a regular brood in the canary. This reproductive trait seems not to be affected in this
cross, which is certainly logical since it is the female canary and not a hybrid female.
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 female 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 main problem
to obtain the red canary was precisely the yellow color contribution from the yellow female
canary. Then, he accomplished the red siskin x dominant white female canary cross, expecting
that the white dominant 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 plumage. Unfortunately, it was not that way
either. The F1 offspring were copper and ash grey color, in both sexes. So, it was confirmed that
introducing the dominant white did not do any better in obtaining a good red, and not a copper
color. From this last experiment it was obvious that the dominant white gene inhibit the expression of the red color as it does with the yellow color.
Dr. Dunckers opinion suddenly changed course; then he stated that it was mandatory to
accomplish the cross red siskin x recessive white female canaries, since these females has no
gene for yellow color. According to his new theory it was expected a 100 % red offspring. This
theory was fully stated in 1929. In 1933 Mr. Gill accomplished this cross, then obtaining all
males copper color after molting, and all females were grey. Not even one single red F1 was
obtained; besides, all copper color males were exactly the same as those obtained during the
crosses red siskin x yellow or dominant white female canary. These F1 copper color from red
siskin x recessive white, when in turn backcrossed to recessive white female canaries, yielded an
offspring (R1) comprising oranges, yellows, and recessive whites.
Then again, all theories were facing a true practical reality that turned them into a doubtful
ground. If the recessive white and the red siskin were yellow free genotypes and according to
Dr. Duncker the copper color was a by-product of red-yellow combinationwhere the yellow
factor came from to yield copper color in F1 offspring, from red siskin x white recessive
female canary?
In fact, Dr. Dunkers suppositions as the red siskin and the recessive white were yellow free
birds, and that the yellow factors mixed with red factor yielded a copper color F1, both were
wrong. This has been evidenced by the practical experience, which has fully enriched the knowledge 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, conclusively 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 pig24

Fundamentals of Color Genetics in Canaries


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 simple to understand that the red plumage phenotype is not only an open expression of its genotype, but also of the environment, too. This is completely true. In genetics it is
clearly stated that:
Phenotype = Genotype + Environment
Today, we enjoy beautiful red color canaries due to a diet where carotenes are present in concentrated quantities, easily absorbed during the metabolic process for those birds genetically
able to transform them into red lipochrome. Nevertheless, up to this moment the red color
present in the red siskin has not been really and completely accomplished 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 males in a small proportion in the offspring, with no possibilities for female fertility, at all. Nowadays, in the light of all gathered knowledge over the years around the world, it is
well known today that F1 males are fertile in a high proportion, and that females are mainly
infertile. Besides, in the R1 cross (F1 x canary) fertility level is a little bit more reduced, much
more in females than in males, since males could be fertile to a larger extent. When R1 males
are backcrossed to female canaries, then a remarkable difference is observed in the offspring
fertility levels, since males and female are fertile, as well; nevertheless, sporadic cases of infertility may 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 problems will be over, since the
whole offspring in normally fertile. The chart below shows what has been explained:

25

Octavio Perez-Beato
Today, canary breeders all over the world use red female canaries to hybridize with the red
siskin in all crossbred levels, according to the previous diagram. Nonetheless, a deeper red color
has not been accomplished, 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
formula, together with a very easy assimilation process, which outcome is an excellent red color
compare 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 most popular theories on the red factor canary,
according to its theoretical genetic formula. A portion of this formula has been previously
explained in the chapter dealing with white canaries and in the chapter about yellow canaries,
all these genes are also common in the red factor canary.
The red factor canary genetically needs to bear the K gene to normally assimilate the carotenoid
substances, plus the G gene in order to transform these substances into yellow pigments. If the
red factor canary would bear the recessive g gene in heterozygosis condition, a proportion of the
brood would be homozygote for that allele, and this is only possible in theory, since the recessive german white canary has never been obtained; it is only and exclusively a theoretical possibility, as has been explained somewhere before in this book. Besides, the red factor canary
possesses the f gene in homozygosis (ff), since the heterozygote condition (Ff) pertains to a
dominant white canary, in which case its red color shall be fairly hidden. Based on all of the
above, we may arrive to the conclusion that the genetic formula for the red factor canary could
be KKGGff. Unfortunately, with these genes we only can obtain a yellow canary. Obviously, we
need to add something else to obtain a red factor canary.
In modern genetics we assume that the most plausible issue is the existence of a locus R with
two alleles, R and r. The R allele is responsible to elaborate a specific enzyme that transforms
the carotenoids into red pigment inherited from the red siskin. On the other hand, the r
allele genetically means, the impossibility to transform carotenoid into red pigment. Then
again, the genetic formula for a red factor canary could be theoretically: KKGGffRR. Besides,
crossbreeding experiments point to a variable expressivity of the trait, since from parents with
excellent red factor expression on the plumage, part of the brood are not so good.
Now, as a by-product, we can better complete the genetic formula of the yellow canary, based
on what was previously discussed: KKGGffrr since the yellow canary is unable to transform
carotenoids into red pigment, as does the red factor canary.
I would like to mention other simple experiment 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 aforementioned 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

Fundamentals of Color Genetics in Canaries


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, nowadays we obtained wonderful red factor canaries based on special addition to canary foods of highly 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 betacarotene 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 extremely deep scarlet color is
determined by several gene pairs, and not by only one. We may discuss about the chromosomal
stabilization that F1 hybrids go through, as well as the subsequent descendant lines R1, R2, etc.,
but that topic is not in the objective frame of this book. I do really believe this topic together
with other biochemical aspects, must be studied for those who are in research work about the
gene complex 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 emerge,
unless a very specific mutation occurs, which is a lot improbable.

The Red Mosaic Canary


This bird emerged after the hybridization between the canary and the red siskin. The latter
shows an apparent dimorphism between both sexes; the female only shows the red color on the
lesser coverts, primary coverts, breast and rump. The dimorphism in the mosaic female canary
is a by-product from the aforementioned hybridization with the red siskin, though the red
lipochrome is not exactly located as in the red siskin female, since the genome of the canary
species probably induced variations to this pattern, to some extent.
Several experiments were conducted by Matern in Germany, Gill in England, and by Kerrison
and Bennett in the United States, regarding the mosaic trait in the female canary. They concluded that by using these mosaic females (formerly: dimorphic), it was possible to obtain a
deeper red color in the canary. It was Gill who, unquestionably, stated that the mosaic 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, lesser 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 aforementioned areas, which usually are a little bit more colored, though it is not always the case.
After the first molt, the new growth shows the typically chalk-white plumage with all red markings well visible, and with an outstanding deep red color. Nonetheless, it is important to say that
the well known mosaic phenotype in not exclusively shown by the female canary. Any canary
breeder knows that mosaic male canaries are a fact, and that they are really beautiful birds.
27

Octavio Perez-Beato
Before their first molt, the male canaries that bear this mosaic trait are not different from any
common lipochrome bird, as discussed in a previous paragraph concerning the mosaic female.
When they complete their first molt and the new growth show up, an increased red color is
apparent almost all over the plumage, turning into a pure white the rest of them, at the same
time that the critical areas, as in the females, are of a deeper red color. After the molt process
has been completed, the male mosaic canary seems very much like any other red canary, but a
closer look will demonstrate that certain areas are uncommonly white, not seen in any frost red
male canary. Those areas are: upper part of the neck, around the vent, and the head. These are
technical visual marks that differentiate the mosaic male canaries, from the common red frost
canary.

Genetic Aspects of the Mosaic Trait


The mosaic phenotype in the canary is the direct consequence of the combined action of a gene
and hormone levels. In the majority of bird species where the female plumage differs from the
male, we must take for granted that an interaction of sex hormones and genetic factors, are
involved. From this concept, the mosaic male canary cannot show the same mosaic phenotype
as the female canary, since the sex hormones are completely different; consequently, the interaction should differ from one another.
Experiments have shown that extirpation of the ovary from a mosaic female canary, lead to a
non- mosaic phenotype after the next molt, since no ovary can segregate the sex hormone necessary to interact with the gene responsible for the mosaic trait. Then, such female will become
a common red factor bird.
When aging, the mosaic female may become a common red factor canary, since the ovary will
not produce the sex hormone; or will, but in an insufficient level.
E. H. Kerrison stated that the mosaic trait is controlled by an autosomic dominant gene, that
he called H. This gene is activated by the sex hormone, to finally produce the mosaic phenotype.
As the mosaic inheritance is due to the action of an autosomic dominant gene, the birds with
genotype HH, and Hh, will show the same phenotype. In fact, that gene has a variable expression, since the technical red markings in the mosaic birds differed from one canary to the other.
Canary breeders are always concerned in selecting the best marked birds for the shows, as well
as for breeding purposes. Regarding the homozygous recessive birds hh, being h the allele
responsible for the absence of dimorphism, no interaction is possible between this allele and the
sex hormone, since the recessive allele h, has no response to the action of the sex hormone. The
homozygous birds hh, will be the common, non-mosaic canaries.
At this point it is convenient to say that Dr. Bennett conclusively stated that it was necessary to
mate red frost birds and red mosaics, and never attempt to mate mosaics to intensive, since the
intensive factor allele I- is an inhibitor for the mosaic expression in the plumage. Rigorously,
the dominant I allele (intensive factor) probably produce an epistatic action on the dominant H
allele, and this is the reason why the mosaic factor is never expressed on intensive birds, no
matter if the canary is homozygous HH- or heterozygous Hh- for the mosaic trait.

28

Fundamentals of Color Genetics in Canaries


Once in a while, some regular variations in mosaic males prevent the identification of such birds
as mosaics. If in doubt if a male canary is really a mosaic bearer, just let it mate to a non-mosaic female. If you obtain any mosaic bird in the brood, then it is a clear demonstration that the
suspect is in fact a mosaic bearer canary.
The mosaic trait is common in melanin birds, in all types as: bronze, brown, agate, pastel, opal,
etc. These birds show, together with melanins, the lipochrome color remarkably deep in the
typical areas mentioned before for mosaic females. I would like to express that it is not only
the red lipochrome which is expressed in the plumage of a mosaic bird; yellow lipochrome
could be also present, instead. Unfortunately, canary breeders in general, are prone to take red
factor breeding, and not yellow lipochrome in their mosaic birds. It is not uncommon to hear
a surprised rooky canary breeder when he/she gets in touch with yellow mosaic canaries. In
my opinion these birds are as good as the red ones, it depends on how we project canary breeding purposes. We must remember that, primarily, a good show is determined by the diversity
of birds registered.
Those canary breeders devoted to the raising of yellow mosaic birds, ought to be very careful in
mating their canaries. A rule of thumb is to select the excellent, among the best, and try not to
go out of the technical procedures recommended in breeding mosaic canaries.
Next, some guidelines and mating examples are shown in order to obtain yellow mosaic canaries.
If any fancier wants to obtain yellow mosaics, and by the moment time only possesses a red
mosaic heterozygous female as the worst-it is possible to mate this female to a yellow frost
non-mosaic male, then:
Yellow frost male
KK GG rr ff ii hh

red mosaic female


KK GG RR ff ii Hh

and

KK GG Rr ff ii Hh (b)

The F1 offspring is:


KK GG Rr ff ii hh (a)

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

F1(b) mosaic female


KK GG Rr ff ii Hh

29

Octavio Perez-Beato
The R1 offspring will comprise 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 male and females yellow mosaic heterozygous canaries, and then
again, phenotypically they are true mosaic canaries. Consequently, we can introduce the mosaic factor in melanin canaries; in short, we will be handling the mosaic factor in both, lipochrome
and melanin lines at the same time.
I think it is convenient to demonstrate that, if the original mosaic female that yielded the F1 offspring, was homozygous for the mosaic factor, instead, the whole F1 brood would be heterozygous for the mosaic trait, and then phenotypically mosaic birds:
Yellow frost male
x
Red mosaic female
KK GG rr ff ii hh
KK GG RR ff ii HH
The F1 offspring is:
KK GG Rr ff ii Hh (mosaic males and females)

Some Important Remarks on the Red Siskin


It was Swainson in 1820 that reported and described for the first time the red siskin, then scientific name was Carduelis cucullatus, today accepted and also known as Spinus cucullatus.
The red siskin inhabits certain territories in the northern part of Venezuela, Trinidad, northeastern part of Colombia, and probably Monos and Gasparee islands to some extent.
Nevertheless, it is extremely 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 comprising a really small wild population.
Facts point to the event that in the 1930s the establishment of the species occurred, due to a
heavy importation of the bird at that time, augmenting 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 centuries, being responsible the Spaniards for keeping a strong grip on this and other finches as well.
Swainsons 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 accomplished the red siskin x canary cross,
at the beginning of the 20th century. Precisely, it was the extreme encouragement for Dr.
30

Fundamentals of Color Genetics in Canaries


Duncker to start in a short period of time his experiments teaming with Reich, ending up with
a red canary in few years.
In the 1940s red siskin populations were dangerously decimated, and this trend continued
through the 1980s. After many efforts, today a Red Siskin Recovery Project is approaching the
goals of conservation. The American Federation of Aviculture, Inc. is involved in this project.
Further information 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 resemble the appearance of the wild canary; that little bird from the Fringillidae family, introduced in Europe about 500 hundred years ago, which
popularity gained the favor of ornithologists and amateurs, as well.
The green canary, according to many specialists, is typically a rustic bird, hence is the variety
that less problems may introduce in breeding and raising procedures. From this green canary,
many mutations have been developed, being the origin of whites, yellows, agate, etc.
In this very chapter, for the first time, we face a melanic canary: the green canary. This bird possesses the two melanins present in the species, it is, pheomelanin and eumelanin; the ground
color is yellow lipochrome; these three pigments make the combination for the green color, as
it is in the wild canary, and inherited by our domestic bird.
Though it is a biological fact, the presence of pheomelanin must not be visible in a good green
canary, which back streaks, wing feathers, and tail feathers, must be black (eumelanin), with no
trace of brown color (pheomelanin).
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 formulas and the locations for genes in sex chromosomes,
from a green male and female. Observe that the Y chromosome is empty, not bearing any gene
of our concern.

32

Fundamentals of Color Genetics in Canaries

Fig. 18 Sex chromosomes carrying the genes for melanin and the oxidative enzyme,
typical of green male and female canaries.

The Brown Canary


This type of canary appeared before 1708 according to some 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 name 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 1908, that two researchers named Durham and Marryat discovered that the
brown color (then known as cinnamon) was the response to a sex link allele due to a mutation.
Besides, this mutation 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 mentioned in previous chapters, published an
article on the color inheritance of the cinnamon tint: Cinnamon Inheritance in Canaries. In this
article the mechanism of inheritance of the cinnamon color (brown color), was explained in full
details.
This brown color on the plumage appeared as a genetic consequence of a mutation, which
inhibits the build-up of black melanin (eumelanin) in the feathers, leading to the solely expression of the pheomelanin, hence the brown color. At the same time, the complete suppression of
eumelanin, 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 definition, a modern tendency led to call them brown, as the accepted adjective to name such
mutation; 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
similar term is brun. Undoubtly, brown is the closest phoneme to these Italian and French
appellatives. Let us say that it is a matter of snobbish terminology, more than a true approach
to the canary color phenotype. It is not the intention of this book to change terminology, but we
need to know why we call a bird the way we do.

33

Octavio Perez-Beato
Genetic Characteristics
To approach the genetic formula of the brown canary, let us call n the allele that denotes the
absence of eumelanin; then we may face the structure of the genetic formula corresponding to
a brown canary, as shown in Fig. 19. The interpretation is exactly like the one on Fig. 18, but
somehow simplified to have it easier.

Fig. 19 Sex chromosomes from a male and a female brown canaries,


with alleles for the expression of pheomelanin and the oxidative enzyme.
The reader should compare Fig. 18 and 19 to fully understand how the brown canary came to
be. Observe that the allele N is dominant to n, which is absolutely necessary to understand the
following steps.
If a brown male mates a brown female, the whole offspring will be brown, as shown in Fig. 20.

Fig. 20 Brown male and brown female mating,


and offspring obtained which is 100% brown.
Likewise, the reader may verify that mating a green male to a green female, the offspring will be
100 % green. Just follow the same procedure as above to test this assertion.
At this point, we are ready to continue setting up couples to mate these melanic canaries. Next
experiment will be mating a green male to a brown female, as shown in Fig. 21. A closer look
will demonstrate that all males in the offspring are carrying the brown factor; on the other hand,
34

Fundamentals of Color Genetics in Canaries


all females are homozygous green, as they only posses one X chromosome cannot carry any
other allele, since the Y chromosome is empty as long as we are concerned.

Fig. 21 Results from a green male and brown female mating.


All males are green carrying the brown factor. All females are green.
What has been stated before is a genetic principle, which is an unconvertible rule regarding
melanin 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 convenient to analyze the opposite mating, it is, brown male and green female. See Fig. 22.

Fig. 22 Results from a brown male and a green female mating. All males are green carrying
the brown factor. All females are brown.
35

Octavio Perez-Beato
In this mating we have obtained all green males carrying the brown factor, as we obtained
before in the previous mating (Fig. 21), but in the present mating all females are brown, not
green as in the previous one.
We can see that females always receive the chromosome X from their father; then again, they
will be pure for the trait included in their fathers X chromosome.
We will continue with another mating: green male carrying brown factor and brown female.
Results are shown in Fig. 23.
Here we have four different offspring types: green males carrying brown as their father,
brown males, green females, and brown females. As the father possesses two types of chromosome X, one carrying the brown factor, and the other carrying the green factor, the female
offspring are composed of brown and green females, depending on what chromosome they
will received, (1) or (2).

Fig. 23 Results from a green male carrying brown factor and a brown female.
Males are green carrying brown factor and brown.
Females are green and brown.
The reader might recall that, the combined effect of melanin and lipochromes was mentioned
in the chapter devoted to such pigments. Precisely, now is the opportunity to talk about the
results of lipochromes, as a ground color for green and brown canaries. Again, these ground
colors could be yellow, white, and red. Besides, keep in mind that regarding the yellow
lipochrome, if carried together with the allele s, the phenotypic effect is lemon yellow, though
in some countries the shows put together all yellow canaries, without distinction. But as I said
before, the allele s is a genetic reality, not a mere speculation. It does exist, and then creates
the lemon yellow.

36

Fundamentals of Color Genetics in Canaries


It is almost mandatory to mention specifically, the varieties that could be obtained according to
the type of lipochrome as a ground color. Such varieties for the green and the brown canary are
listed below:
Lipochrome

Green

Brown

Yellow
Lemon Yellow (allele s)
White (absent lipochrome)
White (plus allele s)
Red

Green (common)
Green (lime green)
Grey
Blue
Bronze

Yellow Brown
Lemon Y. Brown
White Brown
White Brown
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 means can we call it blue. If we
accept this as a fact why not to accept the lemon yellow as a ground color? Then again, we
are not dealing with a subjective criteria, the optic factor expressed by means of the allele s, is
a genetic reality.
Not accepting the presence of the lemon yellow canary is detrimental 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 enzyme for the complete and
full expression of melanins in the green and brown phenotypes, which are both oxidized
melanic forms.
In this chapter, the diluted melanic forms will be the topic. These forms are yielded by an inherited factor that produces an incomplete oxidative process on the melanins, turning them into
diluted forms.
If in the previous chapter the oxidative factor was called O, in the present chapter the dilute factor, or incomplete oxidation will be called o. In this simple way, we are inside the genetics of the
dilute canaries: the Agate and the Isabelle.

The Agate Canary


This type of canary, in some of the varieties, is not exactly a beautiful bird; nevertheless, in other
varieties the combination of colors is really beautiful. This dilute factor appeared for the first
time in the Agate, and it was a turning point for canary breeding at the end of the 19th century.
Not long before 1900, a German canary fancier by the name of Helder raised a female canary of
an unknown color at that time, from a pair of green birds. That female was of a grey tint
plumage, and the lines on the back were reduced, light tinted, and more pale than those of a regular green canary. The first name for this color was ash-grey.
Some 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 1708, mentioned somewhere before in this book. Hervieux called this color Common Agate Serin. From
that on, this variety was called Agate, and not Ash-Grey.
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Fundamentals of Color Genetics in Canaries


Posterior conducted research, demonstrated that the Agate was in fact a Dilute Green, and that
it was a sex link factor. This factor is expressed as the impossibility for the oxidative enzyme to
act at full strength, hence the melanin show itself dilute, since it is not completely oxidized. It
is simple to say that the dilute plumage is the one in which the melanic pigments are pale, due
to the action of the dilute gene o.
As a result, the genetic formula for the Agate is:
Agate male
XBNo
XBNo

Agate female
XBNo
Y

Now compare these formulas to the green canarys 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-oxidative 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 mutation, it showed-up from the Agate, as a result from certain breeding, transferring the non-oxidative factor to the Brown genotype, which turned to be
the Isabelle.
I think it is interesting to explain to the reader, how is the mechanism by which, the gene
that determines the dilute melanin, gene o, is transferred from the Agate to the Brown to
create the Isabelle. This mechanism is understood through the genetic phenomena known
as crossing-over.
Let us suppose that a male with green phenotype is in fact a Brown carrying Agate, according to
the formula X Bno X BnO. If you take a closer look, this genetic formula will yield a phenotype
according to a green canary. Then, in a very specific moment during the meiosis, the homologous chromosomes overlap, each one cut-along in two chromatids. The homologous chromatids
exchange homologous segments, 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 chromosomes of Brown canary carrying Agate genes, at a precise moment during meiosis,
exchange genetic material, creating two new chromosomes. In (a) it is observed a pair of homologous chromosomes of a Green phenotype canary carrying Agate genes. In (b), the crossingover is depicted, showing the exchange of the lower ends of the chromosomes involved, where
alleles O and o, are located. In step (c), the chromosomes are getting apart after exchanging
their segments. In step (d) the chromosomes are set apart already, having each one a new segment in their lower portions. Chromosome 1 is carrying Isabelle genes, since the allele B is present, which determines brown melanin (pheomelanin), plus the allele n which indicates absence
of black melanin, finally the allele o, dilute factor. Consequently, this chromosome will be
responsible for a dilute brown canary, which is nothing but an Isabelle. Chromosome 2 is typical for a green canary. Each one of these two chromosomes will go to a different gamete, leading to an offspring of Isabelle and Green females, besides Brown and Agate females, as well.
These last two phenotypes originate from those original chromosomes not having a crossingover process. Now, it is very clear how an Isabelle female originates from Green male carrying
Agate genes, due to the crossing- over process. Furthermore, 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 demonstrated, they produce four different kinds of gametes, which generate the four classical melanic 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 melanic birds.

The Agate Phenotype


Fig. 25 depicts schematically, the head of an Agate canary, stressing the designs that best characterize this type of bird. The crown shows a striated design that resembles the one on the back,
but miniaturized. The eyebrow is melanin-free, so exhibiting the lipochrome ground color at its
best. The so called whiskers are well visible in a genuine Agate. This design is the direct result
of melanin concentration in the feathers of this area, producing a very strong contrast with the
rest of the feathers that show a dilute melanin. Besides, legs and bill, are pink in the Agate, no
trace of melanin is accepted.
40

Fundamentals of Color Genetics in Canaries

Fig 25 Agate head. Melanin and lipochrome areas are shown.


The classic varieties for the Agate are: White Agate, Yellow Agate and Red Agate. Of course, the
list may continue adding White Agate Ivory, Yellow Agate Ivory, Red Agate Ivory, Red Agate
Mosaic, Red Agate Opal Mosaic, and so forth. The most outstanding White Agates are those carrying the optic factor, which provide a delicate silvery appearance, only detectable with peering
eyes. For the Isabelle, the same varieties are also true; and the same consideration for the optic
factor, as well.
It is well known since long ago, that the best mating is accomplished between Agate and
Isabelle, which produce a better quality offspring, than mating the same type. The misguided
breeding, regarding dilute canaries, may result in an impoverished type in the offspring. It is
highly detrimental in bird shows and judgment of the birds, which hardly qualify in the corresponding category; extreme cases lead to disqualify such birds, resulting in a frustrating feeling
for the breeder, besides an embarrass situation among his/her peers.

41

CHAPTER 9
The Pastel Canary: the super dilution factor
In 1960, a mutation appeared affecting melanins, and mainly the pheomelanins, since the
eumelanin did not suffered any substantial change. More than fifty years had passed without a
mutation affecting the dilute condition in the canary plumage. This new mutation affected the
Isabelle, diluting the tint of the already diluted brown color in these canaries, but brown
melanin was apparent in wing feathers, tail, back, and head. On the back, the lipochromic
ground color was remarkable, different from the common Isabelle.
In the Brown canary, this new mutation became particularly different, in such a way that provided the name to this new mutation in the canary. The ground lipochrome color is delicately
blended with the pheomelanin, just like as it is in a pastel painting, therefore, the name pastel.
The lines or streaks on the back and flanks are almost imperceptible, and the canary shows that
same blending of lipochrome and melanin, all over the plumage.
If we can talk about the Brown Pastel and the Isabelle Pastel, something somehow different
occurs regarding the Green Pastel, Blue Pastel, Bronze Pastel, and Agate Pastel, since the Pastel
Factor does not dilute the black melanin (eumelanin), only the brown melanin (pheomelanin)
is affected. It has been a controversial issue, regarding the existence of true pastel tones in the
aforementioned eumelanic types, ever since. Besides, it is not a simple task to determine 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 same way that the Agate mutation occurred, the Pastel factor is a recessive and sex link
gene, too. At this point, we may consider the genetic formula for the Pastel factor, taking into
account that it is a mutation able to partially inhibit the production of brown melanin, through
a different biochemical reaction step that the one responsible for the appearance of the Agate
canary. Therefore, the non-pastel canaries are considered the normal morph, represented by an
allele P, and then, the Pastel trait might be defined by the allele p, as it is a recessive one. Based
on all of the above, the genetic formula for the Brown Pastel is:
42

Fundamentals of Color Genetics in Canaries


Brown Pastel male
XBnOp
XBnOp

Brown Pastel female


XBnOp
Y

It is apparent that adding the genes for lipochrome colors to the previous genetic formulas, the
classic Pastels will be obtained. What follow are the genetic formulas for those classic Pastel
canaries, most commonly obtained in regular breeding procedures.
Yellow Brown Pastel intensive
X BnOp
KKGGffIi

Yellow Brown Pastel frost


X BnOp
KKGGffii

White Brown Pastel intensive


X BnOp
KKGGFfssIi
X BnOp

White Brown Pastel frost


X BnOp
KKGGFfssii
X BnOp

Yellow Isabelle Pastel intensive


X Bnop
KKGGffIi
X Bnop

Yellow Isabelle Pastel frost


X Bnop
KKGGffii
X Bnop

One important aspect to get the best is the optic factor on the white ground color.
White Isabelle Pastel intensive
X Bnop
X Bnop
KKGGFfssIi

White Isabelle Pastel frost


X Bnop
X Bnop
KKGGFfssii

Note that in all white varieties the alleles ss have been included, since they determine the optic
factor on the white color (either dominant 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 something that
some fanciers do not know, being very concerned why their birds do not get good marks in bird
shows
As in any other sex link factor, the pastel factor carried by a male, will be inherited by its daughters. A straight inference leads to the conclusion that, a heterozygous male for the pastel factor,
will transfer it to half of its daughters, and they will be Pastel. It is now apparent that if a male
carrying the Pastel factor mates any melanic female canary, we will obtain Pastel females, by
means of which we may set up a complete line of this type.
Now the reader is almost ready to conduct the necessary breeding to get Pastel canaries, comprising different varieties.

43

CHAPTER 10
The Opal Canary: the extra dilution factor
Chapter 8 was devoted to a type of canary which recalls the name of a gem: the Agate. The present chapter will deal with another gem name: the Opal. According to certain records, this mutation appeared in 1949 in Germany, probably not well understood at that time. Through the
fifties, it took a long journey, and is exactly during the sixties that became well known and popular, all over the world. It appears that, with the first Opal canary fanciers thought it was merely, and nothing but a more dilute Agate. Then, the exact recognition of a new mutation was not
a simple decision.
Between the Agate and the Opal, we may establish a comparison. The dilute factor that made
possible the Agate type, affected both, the pheomelanin and the eumelanin. The same is true for
the Opal, in which an extreme dilution is accomplished to affect both kinds of melanins, much
more than occurred in the Agate phenotype. The Opal feather is darker on the inferior vane portion; it stands as one remarkable characteristic of the Opal phenotype. Besides, it is stiff, somehow fracturable, and not flexible, mainly in the Green, Blue, and Agate. It is precisely a drawback in the Opal type, poor plumage quality, on the average.
Why that name Opal? In fact, the back of this canary shows certain iridescences when red
lipochrome is present, and in melanic 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 more acute dilution than the observed
in a common Agate, mainly on the back of the bird, on brown and black melanins. 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

Fundamentals of Color Genetics in Canaries


The Opal gene swipe off the melanin from the feather edges, without touching the bill, legs and
under-plumage area, in melanic types. Besides, the Opal factor completely dilute the brown
melanin (pheomelanin) so it disappears from the plumage, then the under-plumage appears
bluish, by the solely presence of eumelanin, extremely diluted. As the pheomelanin is not present, the ground lipochrome shows up, shiny and lustrous.
It is so a strong dilution on the melanins that, the Opal canary with black and brown melanins,
looks like a lipochrome canary, but with the iridescences and bluish tones of the Opal factor,
itself. Only very faint streaks of melanin 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 remain, making 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 common
lipochrome canary, except the duvet, which is black pigmented and partially dilute.
The Opal factor acting on the Agate, swipes off the pheomelanin, completely; nonetheless, the
typical Agate design remains, including: whiskers, back and flank streaks, and the crown
design, but all these are reduced to a minimum expression. They are shortened on half of their
width, and smaller in length.

Genetic Features on the Opal Canary


The dilution factor that created the Opal phenotype is recessive and non-sex linked; it is autosomic recessive. Knowing this, inheritance characteristics can be established; then we are in
good shape to make a forecast from the breeding scheme we plan to make. Let p be the allele
that produces the Opal condition, and P the non-Opal phenotype.
A Green male carrying the Opal factor has the genetic formula:
X BNO
Pp
X BNO
A Green female genetic formula, also carrying the Opal factor, is:
X BNO
Pp
Y
Now, let us number the gametes to establish the possible offspring:
1 X BNO
Pp
mated to:
3 X BNO Pp
2 X BNO
4Y
1 + 3 PP = non-Opal Green male
Pp = Green male carrying Opal factor
pp = Green Opal male

1 + 4 PP = non-Opal green female


Pp = Green female carrying Opal factor
pp = Green Opal female

2 + 3 = 1 + 3 (already explained)

2 + 4 = 1 + 4 (already explained)

45

Octavio Perez-Beato
Now, a simple conclusion is: from this mating we may obtain males and females, as follow: nonOpal, carrier of the Opal factor, and pure Opal canaries. It is important to stress that, as the
Opal factor is not a sex-link gene, it segregates independently of the sex chromosomes, 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 some records point to the year 1950, for the
first time in the history of canary breeding a mutation showed-up that affected the lipochromes,
not the melanins. This new mutation appeared from a couple of yellow Hartzer Roller, from
which all the female offspring showed the new color, as a delicate cream, which enriched a little bit more the already up going genetic background of our feathered little friend, and so opening new possibilities to create color combinations not imaginable, just a short time before.
Ivory, as a color, is well known from the teeth of mammals, mainly from the elephant tusks,
which have been used in works of art for centuries. There are good reasons to name the new
color mutation as Ivory. To better approach this new color in the canary plumage, it is necessary to recall that in the yellow canary, the wing and tail feathers, are not completely colored
with yellow lipochrome, and then some white areas are commonly seen. On the contrary, in the
Ivory canary the creamy color is extended to wing and tail feathers, completely.
In fact, the Ivory factor acts on the lipochromes as a dilute gene, besides extending the pigment
to wing and tail feathers.
The fact that the first two Ivory canaries were females, pointed to the certainty that it was a sex
link factor, as it was confirmed lately, besides being a recessive gene.
In this mutation, there is something new added to all of the above. It was almost a rule that the
former mutations affecting the melanins in the canary also inhibited the trait to some extent.
That is why some authors assessed that the mutation which created the Brown canary, is such
just because inhibited the eumelanin expression in the canary plumage. In addition, the appearance of the Agate is possible just because that particular mutation is able to inhibit the total oxidation process on the melanins. Furthermore, the Isabelle reached on the stage, simply because
of the convergence of two former inhibitory processes: the one that created the Brown canary,
and the incomplete oxidation process that originated the Agate variety.
47

Octavio Perez-Beato
Now, let us examine the Ivory factor at the light of the inhibitory records that, characterized the
mutations on melanins before the appearance of the Ivory mutation. First, we may say yes, this
mutation diluted the lipochrome pigments not allowing deep colors, but at the same time the
wing and tail feathers are now fully lipochromed, not seen before in the canary breeding history. These feathers are so well pigmented 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 same phenomena: dilution of the
lipochrome (inhibitory effect), and extension of the color to wing and tail feathers (antiinhibitory effect). This apparent opposite effects produced by the same mutation, is recorded
for the first time in the canary breeding history.
Advanced canary breeders went beyond the Yellow Ivory. The Red Factor lovers wanted to
experiment 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 plumage. It is a very delicate pink color, evenly extended on all feathers.
In English language it was not a problem to name these two new colors, but in Spanish speaking countries, canary breeders deliberated about a fair nomenclature for both new colors. It was
then agreed that, the Ivory plus yellow lipochrome became Marfil-Crema (Ivory-Cream) according to the exact look of the plumage as a whole. For the red factor Ivory, it was named MarfilRosa (Ivory-Rose) in remembrance of the delicate color of the rose flower. Nowadays, Spanish
nomenclature matches with English equivalent, for a better international understanding of the
always growing color listing, in our modern canaries.

Genetic Features of the Ivory Canary


Now, it is necessary to focus on the genetic formulas 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
X BNOa

KKGGffrrii

Rose (Red Ivory) Frost:

X BNOa
X BNOa

KKGGffRRii

It is evident now, how complex the genetic formulas might be, describing colors in the canary.
Nevertheless, they are necessary by all means to a dedicated canary breeder, who wishes to
completely master in all details, the color genetics on this little bird.
The reader, at this point, must be in good shape to fully interpret the mating procedures that
follow. They will be useful to have a full view of the genetic features of the Ivory factor.

48

Fundamentals of Color Genetics in Canaries


Green carrying Ivory factor male x Green Ivory female
1 X BNOa
2 X BNOA

3 X BNOa
4Y

Offspring:
Green Ivory male
1 + 3 = X BNOa
X BNOa

Green Ivory female


1 + 4 = X BNOa
Y

Green carrying Ivory Factor male


2 + 3 = X BNOA
X BNOa

Green female
2 + 4 = X BNOA
Y

Green carrying Ivory factor male


1 X BNOa
2 X BNOA

Brown Ivory female


3 X BnOa
4Y

Offspring:
Green Ivory carrying Brown factor male
1 + 3 = X BNOa
X BnOa

Green Ivory female


1 + 4 = X BNOa
Y

Green carrying Brown and Ivory factors


2 + 3 = X BNOA
X BnOa

Green female
2 + 4 = X BNOA
Y

The reader surely realized that allele A represents the non-Ivory factor, which is the dominant
original allele. The last mating, depicted in the previous diagram, is the key to understand the
combinations Ivory-Melanin phenotypes, which many canary fanciers believe are somehow
complex to obtain. In fact, it is not. It is just a matter of applying correctly the knowledge
obtained about, dominant and recessive genes, autosomic and sex link, together with the corresponding domain of lipochromes.

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
melanin pigments. Ultimately, the eye color in the canary can provide an extremely important
information regarding melanin 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 may succeed in detecting that important information, needed in the genetics of the pied canary.
Locus P bears two alleles: P and p. The dominant allele P allows the development of melanin in
the plumage, while the recessive allele p inhibits such development. The melanic canary has the
genetic formula PP, a pied canary has the genetic formula Pp. Finally, a lipochrome canary has
the formula pp.
Nonetheless, the spot and shape of melanin patches on the plumage depend upon other genes.
Gene P only allows the melanin to show up on the plumage, nothing else. Now, a question arises: Is there a complete lack of melanin in lipochrome canaries, just because their sex chromosomes do not carry genes for the synthesis of those pigments? The answer is, NO. The
lipochrome canaries do carry melanic traits; their sex chromosomes have the information for
such inheritance. The issue is that a lipochrome canary cannot develop melanic color in its
plumage, since its genotype is pp, which precisely inhibits the development of melanin. A common lipochrome canary may be carrying Green, Brown, Agate or Isabelle. How do we know? It
is not always simple. As an example, a Green, Bronze or Blue canary will show black eyes, since
eumelanin and pheomelanin are deposited in the eyes, as they might be on the plumage. 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 mean albino or lutino, it is not the case. The Isabelle eye color is pink, just in the pupil; albino, lutino and rubino
canarys eyes, are as red as in an albino mouses eyes.
50

Fundamentals of Color Genetics in Canaries


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 eumelanin, and then that pigment cannot be present in their eyes,
either. Based on all of the above, it comes that a lipochrome canary with pink eyes (on light
reflection) is carrying the Isabelle factor; it is not seen on the plumage because the canary does
not carry the gene P, which allows melanin to be seen. In analogous way, a lipochrome 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 some
important couplings. If a breeder has a Yellow male with reddish eyes, and a Green female, of
course, then the following mating is possible:
Yellow male with reddish eyes
X BnO
KKGGffpp
X BnO

mated to:

Green female
XBNO
KKGGffPP
Y

the offspring is as follow:


Yellow-Green pied male
X BnO
KKGGffPp
X BNO

Yellow-Brown pied female


X BnO
KKGGffPp
Y

What is important here is the outcome of the Yellow-Brown pied females, not seen in the parents. The breeder in this example was keenly interested in obtaining birds to start a line of
Brown canaries. The male parent with reddish eyes was unequivocally a Brown carrier, but lacking the allele P to express the melanin in its plumage.
It was then necessary a specific female to provide the offspring with such allele. The chosen
Green female provided the P allele, ending up with an offspring of Yellow-Brown pied females.
From these females, now it is possible to develop a whole line of Brown canaries. To accomplish
this is simple, just selecting the best of the Yellow-Brown pied females and make a back-cross
using its father (inbreeding):
Yellow male with reddish eyes
X BnO
KKGGffpp
X BnO

mated to:

Yellow-Brown pied female


X BnO
KKGGffPp
Y

Offspring:
Yellow-Brown pied male (A)
X BnO
KKGGffPp
X BnO

Yellow male (B)


X BnO
KKGGffpp
X BnO

Yellow-Brown pied male (C)


X BnO
KKGGffPp
X BnO

Yellow-Brown pied female (D)


X BnO
KKGGffPp
Y

51

Octavio Perez-Beato
Yellow female (E)
X BnO
KKGGffpp
Y

Yellow-Brown pied female (F)


X BnO
KKGGffPp
Y

From this entire offspring is now possible to choose from three groups of males, and three of
females. Our choices increased, and then our possibilities to focus on the creation of a Brown
genetic line from the initial mating. It is a good choice to select males from group (C), and
females from group (F) since both groups show homozygote birds for the Brown factor. From
these males and females, it is possible to successfully develop a whole lineage of Brown
canaries. At this point it is advisable to use inbreeding methods. By all means,
there is not a better choice to create lineages as the one discussed in this chapter. The reader
has been provided with inbreeding methods in a previous chapter.
Now, is the time to go directly to the practice of inbreeding methods, together with a strict selection of the best, to ensure a good quality lineage to begin with.
As the reader may see, the most important action is to start with the chosen melanin hidden in
a lipochrome phenotype, focused on the point that the offspring could get the allele P, in order
to express the melanin on the plumage, this allele has been obtained from the non-lipochrome
parent, and introduce in the offspring by means of the right mating system.
In this very chapter only the example of a Yellow carrying the Brown factor, has been discussed,
but it is easily projected to lipochrome canaries carrying Green, Isabelle, etc. depending on the
decision of the canary fancier. The ultimate goal is to obtain birds with phenotype PP, to fully
express the melanin on the plumage.

The Variegated Symmetrical Canaries


According to well documented records, during the XIX century symmetrical variegated canaries
were highly valued, probably due to the fact that it has never been easy to obtain such varieties,
not to mention the hard task in keeping a whole lineage of such canaries.
It seems that, common pied canaries were kept as good pets. Since breeds as the Hartzer Roller
were mostly pied, and the song remarkable sweet. Then, when a more technical approach was
placed on the stage of color canary breeding emergence, 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 symmetrical variegated canary; an
outstanding bird difficult to cultivate and breed successfully, then becoming a highly valued
canary ever since. In this way, variegation lovers were at please, with a new modality of variegated 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 common to find descriptions such as:
brown tail and yellow body, dark crown in a light plumage, etc. Symmetrical variegation is
also a well established feature in Type Canaries, as in the lost London Fancy. It is clear that in
many countries, in one way or the other, the fanciers looking for symmetrical variegated birds,
really abounded.

52

Fundamentals of Color Genetics in Canaries


Dr. Duncker also contributed in the genetics of symmetrical variegation in canaries. He conducted some studies to determine the inheritance of such fashion. I may 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 meager. Based on Dr. Duncker studies the main nomenclature details on these canaries
are known from Germany, and so the terms to name them. Examples to mention are: Sattel,
which means saddle, and Mucken which means mosquito. These German words are applied to
specific designs on symmetrical variegated canaries. It is called point or mark where the
melanin shows up in a regular distribution to create a symmetrical design. In this way, a bird
with melanin around both eyes, and on every tail feather, is said to be a three-point bird. Figures
26 and 27 depict some classical symmetrical variegated canaries; by no means are they a comprehensive picture-list, other combinations are also possible. They represent only schematic
approaches, to give an idea of what a symmetrical melanin distribution might be.

Fig. 26 Upper: Mucken (forehead and throat)


two-points bird. Lower: Also two-points bird,
crown-forehead and tail. Nowadays, color possibilities to create symmetrical patterns in canaries
are really huge, the limit is your imagination and,
of course, the technical possibilities to carry on
mating procedures to achieve what the genetic
projects in our minds are.

Fig. 27 Upper: Typical Sattel. According to


England definition, the symmetrical melanin section is framed on the mid dorsal portion. Lower:
Symmetrical variegated of four points, melanin
patches on both eyes, and on both wings

53

Octavio Perez-Beato
As a general rule, it is recommended to use inbreeding procedures in case we detect an individual with acceptable characteristics of symmetrical variegation. Only using inbreeding techniques, it is possible to achieve certain results in terms of the offspring showing the desired
symmetrical 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 emergence of three new varieties broke all the records known before. Besides, the
new colors are based on the same inheritance pattern: autosomic recessive. They are really new,
so much a new that they are not seen in many bird shows around the world, yet. Perhaps,
Europe is the top of the line leader in modern colors. Nothing new, Europe has ever been. Is it
just because the wild canary was first introduced in the old continent, and their amounts have
been large enough to develop a generous genetic variability? I have no answer, but chances are
that the amount counts, since probability is a matter of large numbers.

The Eumo
In 1981 the Eumo 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. Pheomelanin is almost gone. There
is something that seems 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 terms, the Eumo mutation diminish the amount of melanin; both, in length and
width regarding the streaks. The overall design is somehow streaky, and the non-melanic portion among the streaks is remarkable sparkling, and shows a neat lipochrome color.
The Brown, Isabelle, and Agate Eumo keep their eyes really red, and brightly. The eumelanin is
shown as a very peculiar grey color, including the duvet, but not black. Beak, legs, and nails are
pink.
Eumo is an autosomic recessive mutation. There are some few problems with Eumo phenotype;
if it has too oxidized melanin, the eyes of a non-well trained observer, may judge the Eumo as
a very dilute Topaz.
55

Octavio Perez-Beato
The Onyx
Another autosomic recessive mutation. It showed-up in Spain between 1983 and 1984. This
mutation 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-dominant to the Opal allele, each one located in homologous chromosomes, on the same locus. Then, the Opal is a multi-allelic gene.
The history of the Onyx canary is by no means the same as for the rest of alleles affecting the
melanin. All began more than twenty years ago, when a South American bird of the genus
Spinus hybridized with a Green Opal female canary in captivity. In fact, it is not very clear if it
was a Green or a Bronze Opal female canary; a Blue Opal could be another possibility.
According to what is known today, some hybrid males were fertile. The back-cross to female
canaries, during a certain period of time, came up with the first Onyx canary. As the reader may
see, it is not a mutation at all, but the direct result of hybridization with a species of the same
genus (Spinus) of the red siskin. Records do not specify what the species was. Just for the reader to have a rough idea about what numbers are involved in the genus Spinus, let us say that at
present twenty-one species are recognized with 36 sub-species as well. You may imagine 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 individuals were obtained, and obviously, an improvement in the stabilization of the Onyx phenotype, was achieved.
In fact, the Onyx is the opposite of a dilution. The Onyx plumage is very dark, somehow a blend
of black and brown melanins, with some ash-grey color in the lighter areas. When young, they
look as classic melanin canaries. The appearance of melanin is somehow delayed, but finally
spreads all over the plumage, in a non-evenly distribution. The tail and wing feathers do not
show an even allocation of melanin, on the contrary, it looks like a dash-line design, on a transverse fashion. There is an evident concentration of eumelanin on the head, from neck up, but
this eumelanin does not look exactly pure black. The rest of the plumage shows a disperse
melanin color, more than in any other phenotype known today. Onyx is not a black canary, as
its name may suggest; lipochrome is seen among the streaks, though somehow darker due to
the spread of melanin. The pheomelanin 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 seems reasonable that as the hybridization went on, a crossingover took place; exchange of genetic material between both species the canary and the South
American Spinus-was consummated. Then, a new allele from the Spinus species was added to
the Opal locus. Now, the Opal and the Onyx are co-dominant alleles.
Some examples will help to fully understand how this allelic relation works. Let L, be the normal dominant, 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 normal dominant alleleL, the recessive l responsible for
the Opal, and finally the recessive allele l responsible for the Onyx. Do not forget they are all
autosomic, and l and l are co-dominant.

56

Fundamentals of Color Genetics in Canaries


Now the mating of an Opal male and an Opal-Onyx female 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 intermediate phenotype between both, since the allele l and the allele l are
co-dominant.
Now, the mating of a male carrying the Onyx factor, with a normal female, will be:

Then again, the Punnett squares show that 50% of the offspring will be normal (not carrying
Onyx factor). The other 50% will be phenotypically normal carrying the Onyx factor. Now, from
the 50% carrying the Onyx factor, select the best females to be back-crossed to the original
male, then:

Twenty-five percent of the offspring will be homozygous Onyx (ll). Now, it is very clear you
would obtain Onyx phenotype canaries, in a term of two breeding seasons though you would
only have a male carrying the Onyx factor, if you follow the previous procedures.

The Topaz
According to reliable records, in 1985 a new mutation came into sight, again in Europe. This
time the genetic factor was called Topaz, in accordance to another precious gem. The main fea57

Octavio Perez-Beato
ture of the Topaz is an apparent modification of the eumelanin, which is concentrated bordering the quill, and leaving a wide pearl color contour all around tail, wing and covert feathers.
The pheomelanin is also reduced, but beak, legs and nails exhibit a light brown color, depending 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 symmetrical, neat and well delineated. Eyes are reddish when very young,
and darker when adult. The gene responsible for Topaz mutation is autosomic recessive, as it
was the case in the Eumo 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 autosomic recessive mutation on the melanins came into view. It was
around the year 1994, in Germany. 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 darkening process of melanins. It is apparent that, those areas where lipochromes are seen through
a light cover of melanins (throat, chest and abdomen), as is the case of the classic Greens, Blues,
and Bronzes, in the Kobalt it is not a light cover of melanin, it is a more dense dark patina that
completely changes the appearance of the bird, besides a well streaked throat, chest, and
abdomen are apparent. The beak, legs, and nails are very dark. In general, the streaks are cut,
somehow fragmented in the areas mentioned before, and on the flanks and head. In well selected birds these short streaks look like an irregular dashed-design.
This mutation has been officially accepted during 2006, and in my opinion it may be improved
in the years to come. Now, at the sight of four new colors under the autosomic recessive condition, all affecting the melanin expression and design, a very simple question arises: Are these
four last mutations autosomic recessive just because, the possibilities of another sex link mutation affecting melanins is extremely restricted nowadays, and nothing might be expected in that
direction? The answer is not a simple 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 provision on the sex chromosomes is not enough to enter a new mutational process affecting
melanins, 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 time.

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 common 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 possibilities 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 possibility 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 may bring, regarding new colors. The same is true for a real black canary. It
may show up from hybridization, or from a mutation. What is the most plausible one? At this
point, we need to know what the history is about black canaries in the past. In many instances,
the facts in the past may be the answer to the present. Let us roll the historical facts, and then,
you may draw your very own conclusions. Perhaps, at least a dim light on the possible achievement of a black canary may come true.
According to certain records it appears that, at the time 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 immersed in hybridizing the canary and the Black Siskin. This species belongs
to the same family 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 same color. Its
behavior as a cage bird, is not easy to handle, since it shows sometimes itself as a very nervous
bird, very sensible to aviary or cage conditions and management. Finally, most of them do not
survive in captivity, according to certain fanciers.
From all of the hybridizing attempts, nothing was clearly obtained. Nevertheless, the general
opinion is that the Black Siskin x female canarys hybrid, is fertile, but in a very low proportion.
Around 1930, a canary breeder from Argentina succeeded in obtaining several fertile hybrids
from the aforementioned crossing. It is said, not confirmed, that in the first bird show in that
country, that canary breeder showed the first true black canary, together with several Black
Siskins x female canarys 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 common 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 male being dark green pied, and the female
white with black eyes. That canary was completely black, except the tail and a small patch on
the neck, both being white color; besides, some few brown wing feathers.
Also a mutation gave birth to a black canary; the bird was registered at the Chelmsford Bird
Show. The owner A. J. Spooner declared that he made nothing to obtain such a bird, which
came out of a brood from a green female. As a fledging it was not so dark, but after the first molt
it turned to be black.
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Fundamentals of Color Genetics in Canaries


Other attempts regarding hybridization were performed using the Cuban Bullfinch (Melopyrrha
nigra nigra) a small black finch from Cuba, with a small white bar on the wings. I personally
watched a Cuban Bullfinch and female canary,s hybrid. It was very dark, larger than a bullfinch
but smaller than a canary, having a beak resembling that of the Bullfinch. Around 1957, hybrids
were obtained from such mating. One of those hybrids was registered at a Canary Show in
Havana. The bird had black head and chest, some 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 families that
have been used in hybridization experiments with canaries are: The Jacarina Finch or Blueback grassquit (Volatinia jacarina); Negrito de Bolivia (Carduelis atrata), remember, also classified as Spinus atratus; Alario finch (Serinus alario) belongs to the same 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 experiments with female 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 time, probably a long
time, and a lot of patient work with no dismay. Of course, this is not for a rookie. Only experienced canary breeders may 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
frame in genetic grounds, since the objective of this topic is precisely, to fit out the reader with
all previous knowledge to share.
About hybridizing female canaries with other species, enough said. It is time to consider what a
mutation might be for the real outcome of a black canary, based on the gene systems already
discussed in previous chapters.
The first approach or theory is to consider that a Black Canary must be the opposite of a
Brown Canary, just because the Brown Canary is unable to synthesized eumelanin, and a Black
Canary might be unable to synthesized pheomelanin. Then the first mutation step could be the
change of gene B to a gene b, which means no pheomelanin 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 chromosomes shall be:
Male:

X bNO
X bNO

Female:

X bNO
Y

It is clearly seen that eumelanin is completely oxidized with no presence of pheomelanin. But
this is not enough. It is necessary for the eumelanin to be extended all over the plumage, to really create a black feathered canary. We all know that a good Green canary shows wing and tail
feathers completely black; unfortunately, this spread pigment does not cover the rest of the
bird, just tail and wing feathers.
Now we need another magic touch to completely fulfill the dream of a Black Canary. As I said
in the previous paragraphs, the black color (eumelanin) needs to be spread all over the plumage.
Let us assume that the allele responsible to deposit eumelanin in canary feathers, as we know it
today, is the autosomic dominant allele D. Then, the mutant allele to cover the whole plumage
with eumelanin is d, being an autosomic recessive allele. According to all of the above, two
mutations are necessary to produce a Black canary, which is not very feasible, since a single
mutation is a very sporadic event in genetic history; it is too much to consider two mutations 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 may recall that brown feathers were visible in that individual; this may indicate that
it is not completely necessary to inhibit the Brown allele to produce a Black canary, but a larger amount of eumelanin, instead. If it is correct, then only one single mutation is necessary,
allele d, and nothing else.
As I said before, let D be the regular gene responsible to deposit eumelanin in the regular canary
plumage, then d is the mutant recessive allele which is able to deposit enough eumelanin in all
feathers, to create a Black Canary. Then again, a possible genetic formula for the black canary
might be:
X BNO
dd
X BNO
As the allele d is recessive, this means that a Black canary must be homozygous for that allele.
Besides this allele is not sex link, on the contrary, it is autosomic.
According to historical facts already discussed somewhere in this book, the emergence of black
canaries seems to be associated with a recurrent mutation, since it has been repeated over a certain period of time, as recurrent mutations do. Nonetheless, the outcome of a Black Canary
needs the concomitant occurrence of male and female carrying the recessive autosomic allele d,
if this theory is correct. The expected outcome of such mating shall be 25% black canaries,
either males or females, or a combination of both.
Time and effort will solve the problem of the waited so long bird; time and effort go together
in most of mankind goals, and the Black Canary cannot be different.
62

CHAPTER 15
Obtaining Varieties through Simple Pairing
Taking into account that new color canaries are somehow difficult to obtain, since they are not
widely available, it is the purpose of this chapter to present to the reader some simple pairing
procedures and schemes, 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 complete 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 examples in the following pairings, genetic symbols will be created for the
new varieties to make possible a true understanding of different genetic combinations, according 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 autosomic normal non-Eumo factor, and allele e the autosomic recessive
allele responsible for the Eumo factor. Now, let us suppose that the fancier only possesses one
Eumo male or female canary, as the Eumo is an autosomic factor, it does not matter if we start
with a male or a female canary.
If the fancier has an Agate Eumo (male or female) and the goal is to obtain an Agate Eumo lineage, then simply by pairing this Agate Eumo with a common Agate (or Isabelle, which is a better choice), the starting point of the lineage is achieved.
In the following, simplified genetic formulas will be used; for this
purpose, it will not be considered the ground lipochrome.

63

Octavio Perez-Beato
Agate male carrying
the Eumo factor
1 X BNo
Ee
2 X BNo

Agate female
3 X BNo EE
4Y

1 + 3 = X BNo EE
X BNo
Agate non-Eumo male

or

1 + 3 = X BNo Ee
X BNo
Agate carrying Eumo male

1 + 4 = X BNo EE
Y
Agate non-Eumo female

or

1 + 4 = X BNo Ee
Y
Agate carrying Eumo female

Offspring:

2 + 3 and 2 + 4 yield the same results as the previous ones.


For the next breeding season, the original Agate male carrying the Eumo factor will be paired
with the best female(s) from the 1 + 4 or 2 + 4, groups. Then, what shall be obtained from such
backcross is as follow:
Original Agate male
x
carrying the Eumo factor
1 X BNoEe
2 X BNo

Any female from groups


1 + 4 or 2 + 4
3 X BNo Ee
4Y

Offspring:
1 + 3 = X BNo EE
X BNo
Agate non-Eumo male

or

1 + 3 = X BNo Ee
or
X BNo
Agate carrying Eumo male

1 + 3 = X BNo ee
X BNo
Agate Eumo male

The gamete combination 2 + 3 is identical to 1 + 3.


2 + 4 = X BNo EE
or
Y
Agate non-Eumo female

2 + 4 = X BNo Ee
or
Y
Agate carrying Eumo female

2 + 4 = X BNo ee
Y
Agate Eumo female

It is evident that on the second breeding season, Agate Eumo males and females are both
obtained. From now on, the fancier may establish a whole lineage of Agate Eumo canaries,
always using a line breeding procedure to ensure the purity of descendants and of course, selection ought to be applied, rigorously.
This example is valid for any autosomic recessive inheritance as is the case of the Onyx
(already explained) and the Topaz birds. Next, an example using an autosomic dominant
allele will be discussed. A good choice could be the mosaic canary, which is an autosomic dominant gene.
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Fundamentals of Color Genetics in Canaries


It is relevant to say that when you are looking at a mosaic canary, you cannot tell if it is homozygous or heterozygous for this particular trait, as it is a dominant allele. Then, let be allele h the
non-mosaic normal allele, and allele H the one responsible for the mosaic trait.
Now, suppose you only have a male or female mosaic canary, as this is not a sex link gene, no
matter what the sex of your selected bird is. Let us suppose you have a mosaic female, then:
Example 1:

The offspring is composed exclusively of heterozygous birds (Hh), which is logically expected
from this kind of mating. Then again, nobody can tell they are heterozygous, since their phenotypes are typically mosaic.
Example 2:

The offspring is composed 50 % heterozygous mosaic birds, which look like common mosaic
canaries, and 50 % normal non-mosaic birds. These normal non-mosaic birds in the offspring
show themselves as an irrefutable proof that the parental female is a heterozygous bird, for the
mosaic factor.
Now, the last example is devoted to a sex-link recessive factor to complete a genetic trilogy,
which encompasses the three main categories of genes in the color genetics of canaries. Our
model factor in this example is the Brown canary.
If you want to obtain a lineage of Brown canaries and only possess a male Green canary carrying Brown factor, you may pair this canary to a Green female, which are readily available almost
anywhere; the procedure is as follow:
Green male carrying Brown factor
1 X BNO
2 X BnO
65

mated to:

Green female
3 X BNO
4Y

Octavio Perez-Beato
Offspring:
1 + 3 = X BNO
X BNO
Green male

1 + 4 = X BNO
Y
Green female

2 + 3 = X BnO
X BNO
Green male carrying Brown factor

2 + 4 = X BnO
Y
Brown female

For the next breeding season, a Brown female is already available to be back-crossed to the original Green male carrying Brown factor.
The results are as follow:
Original Green male carrying Brown factor
1 X BNO
2 X BnO

mated to:

Brown female
3 X BnO
4Y

Offspring:
1 + 3 = X BNO
X BnO
Green carrying Brown factor

1 + 4 = X BNO
Y
Green female

2 + 3 = X BnO
X BnO

2 + 4 = X BnO
Y

Group 2 + 3 is composed of Brown males, and group 2 + 4 of Brown females. In a couple of


breeding seasons, Brown males and females have been available. From now on, it is possible to
create a complete lineage of Brown canaries.
Then again, autosomic recessive, autosomic dominant, and sex link recessive alleles represent
the core of color genetics in canary breeding. The previous examples constitute a practical lesson 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 important
aspect cannot be overlooked: the quality of a nice profile and a harmonious shape, as an undeniable complement 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.

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Fundamentals of Color Genetics in Canaries


When one talks about shape, the meaning is straightforward: harmony that must exist in all
parts of the body and plumage contour, in any good color canary. What follows is just a guideline to be considered, when breeding color canaries. These aspects make 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. Some 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, might not be considered to meet the standards of the color canary category.
The beak should be short, cone-shaped, well implanted and its middle line runs along theoretically tangent to the lower edge of the eyes. Head might be rounded, the eyes being the gentle
center of it. The neck must follow a cylindrical contour, harmonious, linking together the head
and the body, following a soft curve line. The chest should be rather short, smoothly rounded to
continue in a soft curve to conform the abdomen, 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 visually proportional to the length of the body. Dorsum must continue in a straight line with the tail.
Wings well tighten to the body, with upper edges slightly touching each other and running parallel on the back, without overlapping at the tip.
The longitudinal body axis must be 45 degrees regarding the horizontal line. Legs well settled,
slightly positioned beyond a right angle. No deformities in fingers or nails. The number of tail
feathers must be 12, and wing feathers 18, as a minimum.

67

CHAPTER 16
Concomitant Factors in the Color Canary: other mutations

The Ino Factor


The main characteristic of the Inos is red eyes, due to an autosomic recessive allele. Individuals
with pheomelanin as the only melanic pigment are called Phaeo-inos; different varieties are
known. Besides, for those non-melanic canaries, if theplumage is white they are named Albinos,
if the lipochrome is yellow they are named Lutino, and if the lipochrome is red, Rubino. It is not
worthless to say that Inos are highly diluted birds; hence eye pigments are gone. There is a particular characteristic in Lutinos; it is a certain light greenish shade, usually all over the plumage.

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 articles written by specialists and true authorities, have been devoted to Satine, all over the world.
This peculiar mutation emerged in Argentina, at the end of the 60s. Based on the mutation
home country name and the fact of red eyes in these birds, the first appellative was Argent-ino.
What was really new in these canaries? Very simple, they have red eyes, with identical phenotype as the Europeans Inos, but Argent-inos, nowadays Satine, are sex link recessive, besides
showing a lustrous patina all over the plumage, which has been ground enough to name them
Satine, a word from French language (Satine: satin-like). Exactly, it was France the country
where they became 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 completely dilute the eumelanin, almost does not touch the pheomelanin. As a consequence, it is sometimes 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

Fundamentals of Color Genetics in Canaries


The Greywing
This variety, as long as I know, came to light from the Bronze Pastel. Nevertheless, Greywing
melanins are diluted, except those on the wing feathers, which are tinted with an ostensible grey
color. In some individuals tail feathers are grey tinted, as well.
The aforementioned mutations appear just like desirable additions to the mutation catalogue
already in existence. It is evident that a meticulous breeding, and a very accurate selection may
create valuable birds from a genetic point of view. Of course, the more factors in a single bird,
the more 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 merely to have
canaries with a lot of factors composing their genotypes, it is completely necessary the harmonious arrangements of those factors, to obtain birds that are phenotypically appealing.
Besides, the more complex color mutations present, the more excellence might be foreseen
regarding the shape of the bird, otherwise the mutation-loaded canary showing a bad conformation, may seem somehow like a patchy deformed individual, since the color combinations
enhance any defect, on the body shape and plumage.
Somewhere in this book, some guidelines were written to offer a basic idea of what a proportional and harmonious body shape might be, in our canary. Read those lines again, it is worthwhile to keep in mind some simple concepts about color and shape.
Another aspect I want to stress is the extremely strict selection when dealing with several mutations at a time, to produce really valuable birds. The slightest defect might become a real problem in breeding seasons to come. 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 hassle 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.
Nomenclature itself is not a hard issue; on the contrary, it clarifies every single detail and subtle features, nowadays so common in color canary breeding. In every science or in any branch
of technology, it is necessary to establish a precise nomenclature that makes communication
run smoothly, to express concepts and technical aspects understandable worldwide. Canary
breeding is not an exception.
Internationally accepted, nomenclature of color canaries is a must, not only involving domestic
bird shows, but all around the world these shows are feasible just because an international way
of naming is respected and accepted, based on technical phenotype characteristics, as well as on
organization ethics.

The Basis of Color Canaries Nomenclature


Three elements are the support of the mainframe of nomenclature, for this particular case of
color canaries. These elements are: Variety, Type, and Category.
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Fundamentals of Color Genetics in Canaries


This simple division allows locating and including all possible color combinations existing at
present, being flexible enough to include future mutations to come. Besides, it is quite simple
for novices to understand and apply.
Variety
This partition includes all lipochromic manifestation on the plumage, such as: Red, Yellow,
Dominant White, Recessive White, Rose, and Yellow Ivory. It is precisely the Variety which
grade the quality of lipochrome distributed on the plumage, or its absence, complementing the
Type from a nominal basis. In lipochrome canaries, nomenclature starts on Variety.
Type
Under this division all melanin-related nouns are included, such as: Green, Brown, Bronze,
Agate, Isabelle, Isabelle Pastel, Green Opal, Agate Opal, and Brown Pastel, just to mention few.
As it is apparent, Type could be single or double. Single Types are assigned to classic melanin:
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 lipochrome, since lipochromes are
assigned to Variety. As a consequence, we cannot talk about Type in strictly lipochrome
canaries. It does not exist in those birds.
Category
The category is the third and last component in color canaries nomenclature structure. At present, only three categories are recognized to describe the feather quality in the color canary; they
are: frost, intensive, and mosaic.
Perhaps, for some canary breeders it may sound weird to include the mosaic condition into the
category, as a feather structure. The only possible explanation that we have at hand is that the
mosaic condition is the opposite of the intensive feather; you must remember what was already
explained in a previous chapter about the opposite relation between intensive and mosaic. It is,
never mate mosaics 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
Red
Yellow
White
Red
Red
-

Type
Brown
Isabelle
Agate
Green
Green Opal

Category
mosaic
intensive
frost
intensive
intensive
frost
frost

Variety
Yellow
White
Yellow
White
Rose

Type
Bronze
Brown Opal
Isabelle Pastel
Brown Pastel
Agate Opal
Bronze Opal

Category
mosaic
intensive
mosaic
intensive
frost
intensive

It is apparent that some individuals are lacking the Variety and others are lacking the Type, but
all bear the Category. The previous examples 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 examples, you will find that under the
Type component those names with two words description, the first name is always the oldest
mutation and the second, the newest one.
Next, some Varieties and Types will be described for the reader, mainly for the new comers to
this activity, to get acquaintance of the peculiarities of the most common color canaries. What
follow is not a comprehensive list, it is just a guide, a grosso modo approach to learn some few
details about the most commonly seen color canaries.

Melanin Types without Dilution


Blue Dominant Intensive
To talk about a Blue Type, the optic factor must be present; the canary will look dull grey, otherwise. No pheomelanin traces on the plumage, but well defined and recognizable eumelanin 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 rump. Beak,
legs and nails are fairly dark, to denote the presence of eumelanin.
Bronze
These canaries must reveal red lipochrome evenly distributed all over the plumage. Black
streaks on the back and flanks must be evident; wing and tail feathers black, with very dark legs,
beak and nails. Any trace of pheomelanin 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 nicely delineated on the whitish ground typical of this variety. The minute design on the crown and
whisker, are readily seen, all of which is superimposed on a very light grey color. The coverts
show the Agate typical design, which is basically formed 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 plumage, flanks and back resemble a tiger-like
design.
Yellow Agate Intensive
In these birds, the yellow lipochrome is the ground color for the dilute melanin. All the streaks
on back, flanks and head must be neat and clean cut. Any trace of pheomelanin is a true demerit, since only dilute eumelanin must be seen, which shows itself as a lead color design. In this
particular variety, the tiger-like appearance is exceedingly shown.
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Fundamentals of Color Genetics in Canaries


Yellow Isabelle Frost
The pheomelanin is diluted in these birds, and only a very light shade of brown is apparent on
the plumage. 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 semi-transparent vane is all
you can see. The yellow ground is utterly seen, mainly on the chest, belly, and rump.

Super Dilute Melanin


White Brown Pastel
The most appealing birds in this variety are those bearing the optic factor in homozygous condition, 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 factor with a ground white color turns the plumage toward a lead-look appearance.
White Isabelle Pastel
The Pastel factor creates a more 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 appearance, much genuine than the lead-resemblance 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 pigmented. The same is true for the Bronze and Blue varieties. Pheomelanin is almost
gone, and eumelanin is severely reduced. Iridescent reflections are seen on the plumage, and
the general appearance is that of a lipochrome individual.
Bronze Opal
On these birds, the typical iridescent reflection of the Opal canary is highly enriched, due to the
red ground color. Regarding melanin dilution, the effect and appearance are the same as
described for the Green Opal.
The description of the aforementioned birds is just an approach, for the reader to have an idea
of the phenotype details that have to be considered, when breeding these common varieties. It
is not recommended to start breeding any variety without knowing exactly what details and features should be considered in those birds. Objectives have to be very clear from the start; undesirable characteristic may show up, otherwise.
In many countries, the nomenclature does not consider the presence of the optic factor. This
means, for them no matter if the white or yellow ground is combined 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 independent of any classificatory criteria, or any checklist convenience.
Nowadays, the amount of mutations already seen in our feathery friend, make possible an
incredible number of color combinations in Type and Variety that show themselves as really
complex. At the same time, all these combinations confer an outstanding genetic value to those
birds bearing them. It is possible to bring into being birds that combine in their genotypes four,
perhaps five of all available mutations known today, together with factors inherited from the
Red Siskin, such as the red color, and the mosaic factor. Just to mention a few of these possible
genotypes, as examples 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 aforementioned canaries are beautiful masterpieces of genetics, obtained
through a very careful and technical breeding, not commonly seen in bird shows; they are rather
scarce. The more genetic factors put together in just one individual, the more accurate breeding
and selection procedures have to be accomplished, simply because the combination of multiple
factors might create a conflict in the phenotype expression for each individual factor, and for a
whole harmonious 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 methods and principles, due
to the particular aspects of canary genetics, being comparable to any other domestic animal
breeding, at a world wide level. I would like to encourage all fanciers involved in color canary
breeding to make a try in obtaining those types and varieties seldom seen around, just because
those birds are beautiful. It is not a secret that many fanciers do not attempt breeding the complex genotypes, due to lack of enough knowledge, mainly on genetics ground. I really hope that
every topic included in all previous chapters, could be a real encouragement to every color
canary breeder, up in the way to better birds at each breeding season.

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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 domestic animals breeding activity. Nonetheless, though outstanding advancements have been achieved in practical management procedures, and reproductive aspects of
canary breeding, just fewprobably very few fanciers all over the world, have been involved in
a numerical evaluation of their breeding results. Honestly, this is the impression I have after
checking published texts, and lots of references and articles in the internet. Most of that information, not to say all, is based on individual practical experiences, which may differ from one
breeder to another. Then a question arises: What are the quantitative results of all these practical advises? To answer this question is dedicated all of the following.

The Quantitative Idea


Quantify is nothing but put results as numerical values; otherwise, only qualitatively shown and
of course, incomplete. The most simple and outstanding advantages of quantifying are:
Concentrate in just few figures the work results of several years
Interpretation simplicity
Precise information without ambiguities, at a single glance.
To take smart decisions according to quantified results obtained.
Nevertheless, some breeders may state that, if quantifying is taken as a procedure, Mathematics
is involved in some way. The answer is, yes. But only very basic Math concepts are going to be
used, and the results are worthwhile based on so simple Math procedures. I do agree that the
method of quantifying, no matter what the application area might be, is not for everybody. A
basic Math knowledge is necessary. For those that are willing to use those basic mathematical
skills, this chapter has been written.

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Octavio Perez-Beato
Quantitative Information
The so called quantitative information is usually envisioned in two basic forms:
1.- Relative figures (usually percents)
2.- Absolute figures
Relative figures are a very popular manner to offer results, being comprehensible for everybody,
since in modern times it is a common issue that TV programs offer a lot of information 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 information is needed. This information is the raw material
of the analysis, and ought to be collected in the most reliable way. It is supposed that the breeder keeps all this information in his/her breeding log. Such information is as follow:
Total females used in breeding
Total nests
Total eggs laid by each female, with dates.
Total chicks hatched
Total chicks full-fledged
Laying range
All of the above could be registered in simple cards designed for such a purpose, or could be
computerized if an ad hoc program is available. No matter what the registration procedures
might be, what is really important is to register the necessary information.
It is unacceptable an aviary without registering the necessary information. 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 mentioned before, so
no ambiguities may arise. At the same time, it is necessary to provide the reader with a standardized method, in order for everyone to apply procedures in the same way. No drifting interpretations could be allowed for a proper understanding of methods, and then results.

Definitions
Total females used in breeding: it is the number of female canaries that took part in the breeding season, constructing their nests, laying eggs (from 3 to 5 eggs per nest; as an exception 6
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Fundamentals of Color Genetics in Canaries


eggs), and then, incubation. If from those incubated eggs hatching was obtained or not, that is
another issue. The point is that the female canary took part in the breeding season.
Total nests: it is the total amount of nests where eggs were laid, and then hatched.
Total eggs laid by each female: it is the total number of eggs laid by each single
female canary during that breeding season. It is mandatory that egg-laying information 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, completely.
Total chicks full-fledged: it must be considered as a primary concept that, every
single fledging set apart from its parent to strive for itself is a full-fledged chick.
Laying range: it is the time interval (days) from the first and last egg laid in that
breeding season. No matter what hen laid the first or the last.
Once all the elements that will be used as raw material in the analysis of the results have been
thoroughly stated, it is time to go directly to the basic formulas for quantitative purposes.

CHICKS PER NEST (Cn):


Cn = total chicks hatched / total nests
This formula tells about the average chicks per nests in each breeding season. It is an indicative
measure of hatching potential for each breeding pair per nest. Theoretically, the value of this
formula (Cn) must be from 3 to 5; being 3 the minimum acceptable value, and 5 the optimum.
Any value near or larger than 4, is considered good. Then again, 5 is optimum.

CHICKS PER HEN (Ch);


Ch = total chicks hatched / total females used in breeding
Ch is an average fertility measure for each hen that took part in the breeding season. What follows is a simple table as a reference for the breeder to compare his/her results with the theoretical expected values. The table comprises from 1 to 4 broods; this last figure is not recommended, since it is detrimental to the health condition of breeding pairs. Results are divided in
three categories: poor, good, and excellent.

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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 fancier is not exactly referred on the table. Let us put an example 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 column indicating 3 broods, the Table value for good results is 12. The figure 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 formulas 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
formulas are fully explained.
For three chicks per nest the Assessed Potential is calculated by:
Pa = [ 33.3 x Cn] 100
For four chicks per nest the Assessed Potential is calculated by:
Pa = [25 x Cn] 100
For five chicks per nest the Assessed Potential is calculated by:
Pa = [20 x Cn] 100
Now, the interpretation and application of this formula will be discussed in detail. First of all,
this formula result is a percentage value. Then, the Assessed Potential is expressed as a percent,
and accurately measures the goal fulfillment of the breeder in obtaining 3, 4, or 5 chicks per
nest. That is why the value of Cn is a crucial element in the formula to determine the value of
Pa. There are three constant values 33.3, 25 and 20 for each alternative in the formula.
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Fundamentals of Color Genetics in Canaries


Whichever the constant is, its value is multiply times the Cn value, and then 100 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 imposed to his/her aviary for that particular breeding season,
regarding the calculation of Pa. The same is true for 4 or 5 chicks per nest. That is why the existence of the three alternatives in the formula.
The interpretation for values of Pa is quite simple. If the value obtained after calculating Pa
equal zero (Pa = 0), this means that the objective has been achieved 100 %. If the result is larger than zero, then indicates that the objective has been over-achieved in that same value. On the
other hand, if the value of Pa is smaller than zero (negative sign -), this indicates that the objective has not been achieved, and the negative value is the measure below the 100 % fulfillment.
It is my advice to use the alternate formula: Pa = [25 x Cn] 100, since the goal of 4 chicks per
nest is feasible and rational, as an expected result, framed into acceptable biological limits.
Anyway, each breeder is completely free to decide what alternate formula 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 development and origin of this formula, what follows
might be interesting:
Pa = 100 [Cn / 4 ( 1) ] = [ 100 Cn / 4] 100 = 25 Cn 100
Being Cn chicks per nest.
I developed the basis of the formula 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 100
The output value from this formula is a percent. The information obtained is understood
straightforward: how many 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 simple
formula directly points to certain possible problems in the aviary. A low S value may indicate
some problems in every day routine handling, such as:
inadequate feeding procedures
health problems in the breeding stock
inefficient hygiene
parasitism
genetic problems (possible mishandled inbreeding)
Next, rational value intervals for S are offered, intended for the breeder to have an idea of the
way results might be interpreted.

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Octavio Perez-Beato

It is convenient to say that one of the major problems that could negatively affect Survivorship
(S), are misguided inbreeding procedures. Inbreeding itself is not a wrong method; on the contrary, breeds of any imaginable animal species have been obtained by rational and well directed inbreeding procedures. What is unsafe is to apply it mistakenly. Chick mortality during the
first days after hatching could be the result of improperly managed inbreeding procedures.

FERTILITY (F):
F = [chicks hatched / total eggs laid] x 100
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 information about the breeding stock,
males and females as well. If a relatively low number of chicks hatch, compare with the amount
of eggs laid, this means that some kind of fertility problem is arising in our breeding stock. But
not only fertility is involved, among other problems that decrease the hatching level, the following could be mentioned:
female canaries not properly incubating
sudden temperature changes
certain toxicity levels in the food supply
male canaries disturbing and bothering the females, 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 problems that really
exist in the aviary.

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Fundamentals of Color Genetics in Canaries


DEVELOPMENT POTENTIAL (Pd):
Pd = [ S x F ] / 100
The formula that expresses the Development Potential (Pd) has two main elements already
explained: Survivorship (S) and Fertility (F). The product of both factors divided by 100 turns
out the value of Pd. This value in terms of a percent, express the relation between chicks fullfledged and total eggs laid in the breeding season. Consequently, the Development Potential
express the performance of the laying potential, that in turn depends on how many full-fledge
chicks are obtained from the total eggs laid.
The Development Potential (Pd), globally expresses how efficient is the management in the
whole aviary, as well as it includes possible problems in chicks development, from hatching to
full-fledged time.
Next, a reference table is offered with Pd values.

As it was stated before, the Development 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 statement is that
Pd includes the Fertility (F) and the Survivorship (S), as structural elements of the formula.

REPRODUCTIVE FREQUENCY (Rf):


Rf = laying range/ total chicks hatched
This formula tells us how frequently chicks are hatching in the aviary. This figure point to the
progress and production level of full-fledged chicks. An example will be explained to better
understand the information that could be obtained using this formula.
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Octavio Perez-Beato
Let us suppose that there are three canary breeders A, B and C. The results they obtained during 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 means that on the average, every 5 days a
chick will hatch, taking into account that for his/her aviary the breeding season
comprised 78 days, measured 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 performance was
achieved by canary breeder C.
It is not an uncommon issue that, for highly developed aviaries the hatching frequency would
be just a few hours, on the average. This means that the values for Rf are in the order of decimal figures, just like Rf = 0.83, and less.

USING THE INDICATORS IN TIME SERIES


Indicators offer information for a particular breeding season, as was seen before, and the cumulative 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 comparative cumulative table is offered for the breeder to post the information collected every year, and then a chronological log could be kept,
which is the basis to establish a comparison of his/her results in a time 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 comparative table time series-oriented.

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Fundamentals of Color Genetics in Canaries


Comparative Table for Annual Indicators

The year where our records start is called base year. This base year will be the time reference
to compare all years to come, from which a very important information will be obtained about
the progress or retreat, for every single indicator on the table. This information 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, mentioned before. Then, the GENERAL FORMULA to obtain index numbers is as follow:
Index = [target year / base year] x 100
As an example, the index numbers for Development Potential (Pd), will be expressed:
Ipd = [target year / base year] x 100
Based on this example, any convenient notation could be used for index numbers applied to any
specific indicator. All index numbers are expressed as percent values. What follows is a good
example using Ipd (Index for Development Potential) in a hypothetical time series:
Years
2003
2004
2005
2006
2007

Pd
94.3
94.8
96.4
98.3
92.5

The year 2003 is considered the base year, according to what was discussed in a previous paragraph. Then, applying the formula for Ipd, the results are as follow:
Ipd = [94.8 /94.3] x 100 = 100.5 for the year 2004. In a similar way Ipd values are calculated
for the rest of the years, always using 2003 as the base year, since it was the first recorded year
for the analysis. Next, are shown the Ipd values according to the applied formula for each year
on the previous table:
Year
2004
2005

Ipd
100.5
102.2

Year
2006
2007

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Ipd
104.2
98.1

Octavio Perez-Beato
It is evident that years 2004, 2005 and 2006 were better than the base year (2003), since the
Ipd values are larger than 100%. On the other hand, the year 2007 shows a decline for Ipd value.
Something happened that year in the aviary, since the trend was increasing in the previous
years, this meant progress. In cases like this, the breeder must carefully analyze what really happened, in order to take smart actions to benefit from.
All of what comprises this chapter, put numerical tools of tremendous practical value in the
breeders hands, as guide lines in the management of the breeding activity, year by year and on
a time series basis, as well. The success of this special task depends on the enthusiasm and
endeavor of the breeder, regarding the numerical aspects in managing his/her canaries. Those
who feel themselves prone to enjoy the simple math 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.

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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 amateur scientists created the first genetically engineered animal. First Edition. Basic Books. USA, 2003.
Chiny, J.: La Formacion 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. Germany, 1924.
Durram, F. M. and Marryat, D. C. C.: Note on the Inheritance of Sex in Canaries. Rep. Evol.
Comm. Roy. Soc. No. 4: 57-60. England, 1908.
Eyckens, J.: Canarini di Colore. Edizioni Zootechniche, Italia, 1963.
Gill, A. K. : Cinnamon 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. Freeman. USA, 2006.
Gottlieb, F. J.: Genetica del Desarrollo. Edit. Alambra S. A. Madrid, 1968.
Hutt, F. B. Genetica Avicola. Salvat Editores, S. A. Barcelona, 1958
85

Hutt, F. B., and Rasmusen, B. A.: Animal Genetics. John Wiley & Sons. USA, 1982.
Lamb, Bernard C. : Applied Genetics of Humans, Animals, Plants and Fungi. Imperial College
Press. England, 2007.
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,
2006.
Perazzo, B.: Canarios Blancos (dominantes y recesivos). Pigmentos y Formulas 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. Germany, 1971.
Vaccari, G.: Canarini di Colore. Edizioni ENCIA. Italia, 1971.

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