It is not all dominant or recessive. The discovery and documentation of inheritance mechanisms in pigeon colours

In 1865 Gregor Mendel presented his 'Experiments on Plant Hybrids'. From the observation of charac­teristic differences in traits and subsequent crosses, backcrosses and splits, he postulated the 'Mende­lian laws' of heredity. The results did not reach pigeon genetics until after 1900.

Validity of Mendel's laws in pigeons

In 1905 Loisel tried without success to trace the check pattern back to Mendel's laws. Probably irritated by the fact that he also had individuals with red basis colour and grizzles in his stock and with the knowledge of that time could not classify the genetic basis of the combinations.

In 1911 Bonhote and Smalley showed the validity of Mendel's laws also in pigeons in the pattern with the dominance of checks over bars. In 1914 Cole showed that the recessive red in tumblers also followed Mendel's rules.

Check pattern x bar pattern results in heterozygous check offspring, which split in the next generation according to Mendelian rules.


Epistatic effects

In 1907, Bateson coined the term epistasis after experiments with the flower colour of peas.  A reces­sive or dominant gene completely or partially suppresses the effect of a non-allelic other gene.

In 1914, Cole showed that black pigeons differed from those with pattern by a hereditary factor. The dominant spread factor has a covering (epistatic) effect over pattern. In 1922, Sarah van Hoosen Jones demonstrates the largely epistatic effect of spread on extensive data when analysing the pattern. 1928 Metzelaar shows the epistatic effect of recessive red over the basic colours

Homozygous Black (Spread) x Blue Bar gives black offspring. When mated with each other, the pattern hidden under black are revealed in some of the offspring. Here the check pattern.


Sex-linked inheritance

In 1908, Doncaster demonstrated the concept of sex-linked inheritance for the first time in animals using variants of the gooseberry peeper.

In 1911 Bonhote and Smalley and finally Cole in 1912 demonstrated that the mechanism of sex-linked inheritance also applied in pigeons to the mating of blues and the dilute colour silver (dilute blue).  Thus, the phenomenon that silvers from blue parents were always females - which had already puzzled Darwin - had received a resolution. In 1912 Cole showed that this hereditary mechanism also applied to the mating of blacks with dilute dun-coloureds and of reds with dilute yellows. The difference between dominant and recessive red, was documented together with Kelly in 1919. Here they also showed the sex-linkage of dominant red.

Homozygous dilute cocks (here yellows) x non-dilute hemizygous females (here reds) result in hemizygous yellow females and heterozygous red cocks


The inheritance of Pigeons with the genes reduced, Carl Graefe 1951, and Rubella, Gerhard Knopf 2002, follow the same rule.

Rubella-cock x non-rubella hen (upper line) results in cocks heterozygous rubella, and hemizygous rubella hens.


Hereditary sexual colour dimorphism in pigeons

1868 Darwin discusses the possibilities of creating strains with colour differences in cocks and hens. As an example, he finds wine-red pouters in the literature, in which only the cocks would show black spots. Not explainable at that time, according to the analyses of Hawkins (1931) they were heterozygous cocks for black and dominant red basic colour.

In 1925, the Norwegians Christie and Wriedt analysed the dominant sex-linked stipper gene in Danish Tumblers. Indirectly they show the characteristic whitish colouring of homozygous cocks.

Homozygous stipper pigeons are whitish in colour and usu­ally have health deficits in contrast to hemizygous females (here a pair from the author's breeding).


1942 Hollander shows hereditary sexual dimorphism in faded with the possibility of creating reproduc­ible auto-sex strains (Texan Pioneers)

Sexual dimorphism in ash red faded Texans. The lighter-coloured homozygous cocks re­produce when mating with hemizygous frosty females


1970s: TIM KVIDERA discovers in the USA at Racing Homers a gene 'Frosty' probably mutated from Faded. Pure-bred cocks have more distant resemblance to faded heterozygous cocks and hemizygous faded hens on a blue color base. Female pigeons corresponded to the wild type blue-bar. In 1988 he estimated them to be a recessive sex-linked allele of Stipper and Faded. In 2000 ANDREAS LEIß shows that Thuringian Selfs have the same hereditary behaviour. The females, corresponding to the blue-ground-coloured cocks, are blue-bar. After outcrossing upon other breeds there are occasionally slight lightenings in front of the tail band. This is also found in heterozygous faded cocks and is described by BECHSTEIN 1807 in the historical 'tail pigeon' native to Thuringia as giving its name. Unlike Reduced, Rubella, the dilution factors, and also Faded, the hemizygous females in Frosty do not show their sex-linked factor.


Thuringian blue-ground-colored cock and a black non-frosty hen with black and blue bar offspring. At the right a heterozygous cock and a hemizygous frosty hen.


Genetic Linkages

1911 to 1929 Morgan and his co-workers make extensive Drosophila mappings of chromosomes and document frequencies of linkage breaks in genes located together on a chromosome

1919 Cole and Kelly empirically determine a cross-over rate (frequency of linkage breaks) for the sex-linked primary colours and dilution factor of 40% (50% would be the statistical expectation for inde­pendently inheriting genes)

Cross-over test by a cock heterozygous for dominant red and dilute on the same chromosome to a hemizygous dilute coloured dominant red (thus yellow) hen. Source: Sell, Taubenzucht 2019


1938 Hollander determines a close linkage with a cross-over rate of 2-3% for the non-sex-linked factors recessive opal and pattern.

Completion of the basic colours

In 1924 Metzelaar (quoted from Steele 1931) was the first to recognise brown as an independent basic colour. That brown is an allele of Dominant Red was shown by Hawkinson 1931.

Brown as sex-linked basic colour. Homozygous male and hemizygous female do not differ in col­our.


Additional non-allelic interactions

Following Bateson, further non-allelic factor interactions are shown after 1907, especially in plants. Brief introductions easily give the impression that each trait is caused by one or very few genes. However, almost all traits are likely to be the result of cumulative or complementary interaction of several genes (Bartelmess 1965: 724). Pigeon fanciers express this by referring to 'modifying' factors as an explanation for variations judged to be small. In the case of larger variations, potentially significant factors are explicitly named.

1925: That dominant genes override the effect of non-allelic genes is not surprising for pigeon fanciers. Stipper, for example, transforms pure recessive reds in English Short Faced Tumblers and Danish Tumblers into DeRoy. This was shown for Danish Tumblers by Christie and Wriedt in 1925. Fulton (1876) could not yet classify the colouration in English Short Faced Tumblers.


DeRoy English Short Faced Tumbler (left). Youngsters DeRoy, Brown Stipper, Kite and red Agate at Danish Tumblers in the own loft



1929 Christie and Wriedt identify 'bleaching factors'. They only appear on recessive reds that are bred from heterozygous Brander couples. Recessive red thus is the prerequisite to show that effect. The bleaching (whitening) increases with age.


Bleaching type I at Christie and Wriedt, Reces­sive Reds from Danish Brander. Source: Chris­tie and Wriedt 1929


In 1876, Fulton had described recessive red pigeon that moult in parts to white as a complementary colour of the Almond breed. The trait has been transferred to the Danish Stippers bred from English Short Faced Tumblers. Here, too, the gene is only found in recessive reds. In larger experiments Tim Kvidera (1982) for red Viennese White Shields, and Andreas Leiß (2008) for red white shields and Uzbek red Chinny, have confirmed this. They postulated a white-shield factor, which presupposes homozygous recessive red.


English Short Faced Tumbler red Agate, Viennese White Shields recessive red, Chinny Uzbek Flying Tumbler


2010, 2011 Hein van Grouw and Dina Mergeani found the basic ash red red colour to be a prerequisite for the naked-neckedness of Romanian Tumblers. This links a feature to a non-colouring trait.


Romanian Naked-neck Tumbler  (photo at the right: Layne Gardner)


Heterozygous recessive red has been documented as a modifier to improve standard colouration for show in breeding almond English Short Faced Tumblers since Fulton (1876) and later in Danish Stippers. A positive effect on the desired standard colouration may explain why the recessive factor persists in the breed and why recessive red keeps splitting out.


Danish Brown Stipper Cock and Red Agate from Danish Stippers in nest plumage and shortly before completion of moult from the author’s loft. ‘Brown’ is a term of the standard, genetically the basic color is black.

In Copper and Gold Gimpel pigeons, recessive red has a positive effect on the desired copper or gold colouring when heterozygous, analogous to the effect in almonds.  From well coloured parents self reds and golds split out (so already Goodall 1899).


Gold Blackwing Gimpel, heterozygous recessive red, and Self gold cock (Recessive Red and Pale) from Gold Gimpel Pigeons from the own loft Source, Sell, Genetics of Pigeon Colouring (German lan­guage), Achim 2015

1925 Christie and Wriedt found that recessive reds regularly are raised from Danish Brander that moult white-pied (see above). However, they did not connect this observation to bronze colouration of Brander.  2008, 2010 and 2011 Ko van Vliet and others for Dutch Chimney Sweepers, Wim Halsema and Bill Peterson for Danish Brander show that the existence of heterozygous recessive red leads to the trans­formation of otherwise kites into 'Brander bronze'.


Danish Brander and Dutch Chimney Sweeper, heterozygous for Recessive Red

2012 Interaction of recessive sex-linked genes: Frosty and Rubella. Both factors are located on the sex chromosome (see above). One difference is that in Frosty the factor is only shown by homozygous cocks. In Rubella, the hemizygous females also show it. Based on Rubella, the Frosty factor, which does not show in heterozygous cocks and hemizygous hens, has a colour effect. Hemizygous Frosty-Rubella females turn ice-grey in body plumage. Checks and bars are weakened in colour. This was also shown by heterozygous cocks when mating Frosty-Rubella hens with the wild type. Often less distinct. Pattern are still slightly brownish or only anthracite toned down. Pure-bred Frosty-Rubella cocks are light silver-grey. In colour contrast the family corresponds to sex-dimorph coloured faded. Due to the relatively long distance of the loci for Frosty and Rubella on the sex chromosome, the connection is fragile. It is quickly lost in outcrosses.



Homozygous Frosty-Rubella-couple with distinct sex dimor­phism (left), heterozygous frosty-rubella cock from wild-type x frosty-rubella hen (right) at the author’s loft.


Molecular genetic anchoring of genes

In 2004, Charles Lee, in collaboration with a Canadian research group, published a study in the context of the analysis of the human genome in which it was shown that larger sections of DNA were present in all individuals with different copy numbers (Copy Number Variation CNV). In the analysis of the hu­man genome, potential connections with diseases are of particular interest.

2017, 2019, 2020 Domyan, Shapiro, Bruders et al. present extensive studies on the anchoring of genes in the genome and investigate differences and effects, e.g., of CNV for Stipper and alleles of the Stipper-gene.

Copy number variation at Stipper and Alleles of the Stippergene. Source: Excerpt from Bruders R, Van Hollebeke H, Osborne EJ, Kronenberg Z, Maclary E, Yandell M, et al. (2020) A copy num­ber variant is associated with a spectrum of pig­mentation patterns in the rock pigeon (Columba livia). PLoS Genet 16(5): e1008274.


n=number of tested individuals.


Literature und Hints on Literature

Hollander, W.F., Origins and Excursions in Pigeon Genetics, Burrton, Kansas 1983

Levi, W.M., The Pigeon, Sumter South Carolina 1941, revised and reprinted 1969

Sell, Pigeon Genetics. Applied Genetics in the Domestic Pigeon 528 p., Achim 2012

Sell, Genetik der Taubenfärbungen (German language), 384 p., Achim 2015

Sell, Critical Issues in Pigeon Breeding. What we know and what we believe to know, Parts I-VI Achim, 60 p., 2020/2021

Introductions in Genetics in English, French and Dutch

Sell, Introduction to Heredity in Pigeons, 80 p., Achim 2022

Sell, Inleiding tot de erfelijkheid bij duiven, 80 p., Achim 2022

Sell, Introduction à l'hérédité chez les pigeons, 80 p., Achim 2022