All about Inheritance of Bengalese Finch Varieties
Results of approximately 15 years of my research are given in this table.
Varieties Mode of Inheritance
Gene Symbols
Wild type
Mutant

Autosomal
Albino, European Recessive
Ae
ae
Albino, Japanese*1 Incomplete-dominant
aj
Aj
Dilute Recessive
Di
di
Clearwing*2 Recessive
Cw
cw
Gray Recessive
Gr
gr
Dominant Pied(Variegated or Mottled)*3 Dominant
pd
Pd
Recessive Pied(Marked White)*3 Recessive
Pr
pr
White feathers on forehead, chin & flights
-
determining*4
Dominant
wd
Wd
White feathers on forehead, chin & flights
-
initiating*4
Dominant
wi
Wi
Bonten (= Crest)(Chunagon-initiating)*5 Dominant (with recessive lethal?)
bo
Bo
Chunagon-determining*5 Dominant
cd
Cd
Chiyoda*6 Dominant with recessive lethal
ci
Ci
Neck frill*7 Dominant
nf
Nf

Autosomal multiple allelic series
Chestnut*8 Recessive to C, Dominant to Cfa
C
Cch
Fawn*8 Recessive to C and Cch
C
Cfa

Sex-linked
Ino Recessive
In
in
Pearl*9 Recessive
Pe
pe

Notes :
 *1. The heterozygote Aj /aj shows pale or a pastel color similar to the Dilute. I call it "the Light color".
 *2. I have two observations on the genetics of the Clearwing. One is the possibility of the cw gene linked to the di gene. The other is that the cw gene may be allelic to the di gene.
 *3. I conceive that the phenotype of Self White is the interaction of two genes : the Pd and pr genes.
 *4. I consider that the heritage of this characteristic is based on the "Initiator Theory on Crest of Budgerigar". This phenotype is a demonstration of the interaction of two genes : the determining gene and the initiating gene. The genotypes of the birds that have some white feathers on the forehead, the chin or the flights(at least on one of these parts) are Wd /Wd Wi /Wi, Wd /Wd Wi /wi, Wd /wd Wi /Wi, and Wd /wd Wi /wi.
 *5-1. I have never obtained the homozygous (double factor) Bonten bird. It is probably because the Bo gene is recessive lethal.
 *5-2. I consider that the heritage of Chunagon is also based on the "Initiator Theory on Crest of Budgerigar". "Kawabata's Chunagon theory" is the following : the phenotype of Chunagon is a demonstration of the interaction of at least two genes : the Bo gene that is Chunagon-initiating and the Cd gene that is Chunagon-determining. The bird that has the Bo and Cd genes indicates Chunagon. The bird that has the Cd gene but does not have the Bo gene is not Chunagon but otherwise visually normal.
 *6. Expression of Chiyoda is variable. Expression of the Ci gene is probably taken with high penetrance. It is also highly probable that some multiple alleles exist. The homozygous (double factor) Chiyoda bird is generally not able to live, dying in the early embryo stage. It rarely survives and becomes a full frilled bird (e.g., King).
 *7. The phenotype combining the Bo gene with the Nf gene is similar to Chunagon that has the Bo and Cd genes. It is visually difficult to distinguish between them. The effect of the Nf gene only appears in the nestfeather stage, and it then outgrows. But in the case of combining the Nf and Cd
genes, the neck frilled feathers do not disappear as an adult (e.g., my "Wa-chiyoda"). I conceive that Chunagon that has well developed frilled neck feathers is a result of the interaction of three genes : the Bo, Cd and Nf genes.   
 *8. The tri-color is a kind of Pied with the di-color. The phenotype of the di-color is a manifestation of two different color patterns that are the black brown-the fawn, the black brown-the chestnut and the chestnut-the fawn. I consider that the characteristic of the di-color is genetically "trisomic aneuploid" because it rarely happens that this characteristic is inherited. Based on my suggestion the genetic structure on the Ch-lous of the di-colored bird is some one of the following : Ch / fa / fa, Ch / ch / ch, Ch / ch / fa and ch / fa / fa .
 *9. There are rarely cases where the pe gene takes on incomplete-dominant. Based on the observation of Mr. Takahiro Yokouchi who is the well known breeder of Munias and Mannikins some of the heterozygous birds indicated pale pearl in the strains mating with Tri-colored Munia or Moluccan Munia (e.g., the BC1 bird : Tri-colored Munia x (Pearl Bengalese Finch x Tri-coloed Munia)).

 Acknowledgement : I thank Ms. Io Munakata for many helpful suggestions on writing in English.

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