The check pattern in domestic pigeons

For DARWIN, the blue-bar rock pigeon was the starting point of the domesticated domestic pigeon. Later colourings would have developed from blue-bar pigeons in the course of domestication. This is also the statement of the ornithologist BECHSTEIN, who kept pigeons himself. He believes to have observed the appearance of the check pattern, along with other colour changes, in semi-wild pigeons in Thuringia (BECHSTEIN 1795). In contrast, CHARLES OTIS WHITMAN (1842-1910) placed the check pattern at the beginning. For him, the bar pattern of the rock pigeon resulted from a 'gradual progressive modification' in the evolutionary process. He points out the similarity of the check pattern in the wild pigeon species, which is particularly evident in their juvenile plumage. The development from check to bar would also be embryonic development. In the mating of checkered domestic pigeons, descended from the rock pigeon, over the generations, the checks would be reduced to 4, 3, 2, 1 and 0 bars, while in his experiments over 8 years the reverse path, from bars to checkered birds, had remained unsuccessful (WHITMAN 1916, p. 16ff., 162). From today's genetic point of view, however, the pathway observed by Whitman in the experiment, from check to bar, is not unusual if the initial population included a heterozygous-breed individual. A more recent molecular genetic study speculates that the check pattern was transferred from the Guinea pigeon to the domestic pigeon after separation of the species (an introgression). According to one estimate, this was only 429-857 years ago (VICKREY et al. 2018). It is therefore interesting to follow the distribution and documentation of checkered pigeons in the literature.

Fig. 1: Inheritance of checker. Source: Critical Issues Part VII

Dissemination and documentation of the check pattern

In today's city pigeons and when domestic pigeons mix with rock pigeon populations, pigeons with the check pattern make up a large proportion worldwide. Among Viennese city pigeons, HAAG-WACKERNAGEL/HEEB/LEISS (2006) found a proportion of checks (31.7%) and dark-checks (24.8%) compared to 37.3% that were barred. 4% were barless. Due to epistatic effects (e.g. the spread factor covering pattern), 9.9% could not be assessed. When observing feral domestic pigeons in Kansas in 1984/85, JOHNSTON/JOHNSON (1989) found 37% barred pigeons, compared to 22% checks and 41% dark checks. In Bangladesh wild pigeon populations mixed with domestic pigeons, there were, among other colors, 11% blue-checks and 75% blue-bars (KABIR 2016).

In the early days of organized pigeon breeding and in the first monographs on pigeons, checkered pigeons had no importance among pigeon breeders. They were not highly regarded, with the exception of some factor combinations with bronze or white in the check outlines. BECHSTEIN (1795/1807) describes it as a variant of field pigeons. In the first extensively illustrated German-language domestic pigeon book by NEUMEISTER (1837), there is not a checkered example among the 123 pigeons shown on the plates. However, the gene for checks may have been present in the so-called white blaze with whitish or bronze wing shields. In the text, the drawing contour of the checks appears under the 'meliert' pigeons listed after the field pigeons in the text, based on BECHSTEIN. Among these also mentioned are larks and laced ones. The former were probably preforms of lark-pigeons, the latter of the later scaled (laced) lynx pigeons. BREHM (1857, p. 92) describes 'carp-scaled' and 'hammer-faced' pigeons among the field pigeons with which they were associated but formed a minority. PETER PAILLOU had already presented an early drawing of a Parisian Pouter with similar scales in England in 1744. Similar to the Pigeon Maillés created from pouters in BOITARD/CORBIÉ 1824 (p. 179ff.) as preforms of the later Cauchois.


Fig. 2: Paris Pouter with the check pattern by Peter Pailou 1744. Fig. 3: Color Pigeons at Neumeister 1837

However, drawings of blue checks existed even earlier. If you go back 429 years, the lower limit for potential introgression mentioned above, then you are in the time of MARCUS Zum LAMM (1544-1606). In Freiburg (Germany) he had pictures painted for his Thesaurus Picturarum and put together a collection. Also mentioned in his notes is a 'Visch Schüppichte or hammerschlegichte Daub'. This was apparently a common name for checks in the region. One of his pictures that has been preserved is a drawing of a blue-check field pigeon. There must therefore have been checkered domestic pigeons long before this time, as they were already common terms at the time. If we go back 857 years, we are shortly before the time of EMPEROR FREDERICK II (1194-1250), who wrote his Falcon Book between 1241-1248. It was completed by his son. In addition to turtle doves, the numerous miniatures also contain two domestic pigeons, which can be classified as checkered when compared to the blue-bars shown. Even further back, the restored murals in Akhenaten's North Palace in Amarna (around 1350 BC) in Egypt show, among other things, a blue pigeon, which HAAG-WACKERNAGEL (1998, p. 46f.) classifies as checkered. Overall, the evidence from the past gives the impression that checks in domestic pigeons occurred mutatively in different places at different times, as BECHSTEIN describes it.

Fig. 4: Blue check field pigeon, Marcus Zum Lamm about 1600. Source: Kinzelbach/ Hölzinger 2000. Fig. 5: Blue check pigeon in the North Palace of Echnaton in Armarna, Egypt about 1350 BC. Source: Haag Wackernagel 1998


Fig. 6: Checkered Dovecot Pigeon (Albin 1735). Fig. 7: Checkered Dovecot Pigeon (Dixon 1851)

Good conditions were found for checks, kept semi-wild, apparently at the time of ALBIN, who drew a checkered pigeon in 1735 as a representative of the 'Dove House' pigeons in England. A good 100 years later, DIXON describes her as a typical inmate of the English 'Dovecots' (DIXON 1851, p. 162ff.). According to DIXON, the checkered variant had already successfully established itself under railway overpasses in London and other parts of England. This suggests advantages for the checks in the fight for survival as urbanization increases.

The increase in reputation and thus the spread in hobby breeding will be linked to the spread of the Belgian racing pigeon after 1800. In connection with this, from the second half of the 19th century onwards, the breeding of racing homer-related fancy breeds such as the Show Antwerp, Show Homer, Show Racer, the German Beauty Homer, etc. On the 50 color plates in the magnificent work by FULTON (1876) there are two color plates with checkered homing pigeons (in England called 'Flying Antwerp' because of the imports via the port of Antwerp form Belgium), and Show Antwerp. On the other panels there is only one checkered one, namely a blue check shield owl. In combination with white scales, the gene for checks can also be seen in pictures of Oriental Owls, Hyazinth, Starlings and Ice Pigeons.


Fig. 8: Show Antwerp and Flying Antwerp (Belgian Racing Homer). Fig. 9: Oriental Owls with the bar and with the check pattern. Fig. 10: Hyazinth Pigeon and Starling with the check pattern. Source: Fulton 1876

Hybrids with Guinea pigeons

WHITMAN had already pointed out the similarity of the checkering of the Guinea pigeon with the typical light triangular spot at the end of the feather in the shield with the checkered of some, but not all, domestic pigeons. A similar checkering can be found in many wild pigeon species. In addition to the widespread turtle dove, whose checks WHITMAN considers to be the archaic form of checks in the pigeon family (p. 50), the special checkering of the Guinea pigeon can also be found, among others, in Columba maculosa and C. albipinnis (p. 163).

Hybrids of Guinea pigeons with domestic pigeons and reverse matings were analyzed in blood group studies of pigeon species. So, in 1936 by IRWIN, COLE, GORDON as well as MILLER/BRYON 1953 and LABAR/IRWIN 1967. Among others, five different antigenic substances were identified as putative inheritance units that were specific for the Guinea pigeon (MILLER/BRYON 1953, p. 407). Hybrids with barred domestic pigeons showed the check pattern and also the loyalty of the racing pigeons to the location of the own dovecote (COLE CREEK 2019). Problems in maintaining and raising hybrids and early mortality are reported (see also GRAY 1958). Some of the hybrids are viable and capable of reproduction, even when mated back to domestic pigeons.

Fig. 11 and 12: Guinea pigeon and Hybrid-hen, lost by illness. Held and flown with the Racing Homer team. Source: Cole Creek at Facebook. Fig. 13: Columba maculosa, Lip Kee Yap, CC BY-SA 2.0 <https://creativecommons.org/licenses/by-sa/2.0>, via Wikimedia Commons.

Diffusion of mutation and introduction of genes through hybrids

Similar behavior, the possibility of obtaining fertile hybrids, and the visual similarity to the checks of the domestic pigeon led to speculation that the pattern was transferred from the Guinea pigeon to domestic pigeons. In view of the sparse references specifically to the checkered pattern in domestic pigeons from the past, this is not an unfounded hypothesis. Mathematical model calculations show how quickly superior genes, whether created through mutations in a population or added through crossing, can prevail when partners are chosen at random. A dominant factor will reach a proportion of around 50% after 400 years with a constant population and number of offspring (per parent) of 5, an advantage of the carriers of the new trait of 1/100 and an initial frequency of the new gene of 1/1000 (Mittmann quoted from KÜHN 1961, p. 250). Random mating can be assumed in feral domestic pigeons and, with regard to coloring, in racing pigeons that are primarily selected for performance. For other breeds, breeders' different selection criteria play a role. In breeding groups in which new factors have been introduced, the factor will combine with other existing ones over the generations. Let's assume that in a population of pigeons, Smoky is present alongside the wild type. Then after a short time the proportion of pigeons with the smoky factor should not differ between checkered and barred pigeons. Close genetic linkages that could hinder mixing over centuries are unlikely. Regionally, there will be differences in genetic makeup in subpopulations due to spatial distances, other environmental conditions, factor interactions that are not directly recognizable, threats from birds of prey, etc. (SANTOS et al. 2015). Such differences have also been found between feral domestic pigeon populations in the megacities of the northeastern United States from Boston to Washington (CARLEN et al. 2021). For example, in tests in which the proportions of certain factor combinations are important, it will make a difference if, for example, barred individuals are chosen from a different region or from different breeds than checkered individuals.

In the case of introgression, the main difference to mutation within the species is that, along with the gene under consideration, the hybrids are heterozygous for other hereditary factors. Some of them were not present in the host population until then. Most of them are probably not visible externally. They could be immunities, genes that influence energy metabolism, etc. They could be species-specific and, in the species that transmits the gene, have emerged mutatively after establishing as a separate species. If transmitted to domestic pigeons, many factors will disappear quickly, but others will become established and possibly expand significantly. The diffusion is likely to start from the point of origin, similar to a mutation, and spread regionally over time. However, under human care, distances do not play a role for newly discovered mutations, as the Reduced example shows (SELL 2012, 2021). In subpopulations into which these factors have penetrated, they will freely combine with each other and with those present, as in the spread of mutations.

Measuring introgression

ABBA-BABA tests: In evolutionary research, potential introgression is usually analyzed using ABBA-BABA tests, in which four populations are compared with each other (DURAND et al., 2011). There are two currently existing populations P1 and P2. Then an evolutionary older population P3 and a fourth outgroup population O. This is more distantly connected to these three ingroup populations. Their genome serves as a reference; their gene expression at the compared loci is designated A. The alternative that P3 has is called the 'derived allele' and symbolized by B. The null hypothesis for the empirical test is that P1 and P2 diverged from a common ancestral population that had separated from the ancestral population of P3 at an earlier time. After P1 and P2 split off from the ancestors of P3, there was no gene flow from any of the groups with P3. The alternative hypothesis is that P3 exchanged genes with P1 or P2 after these two populations separated (DURAND et al. 2011, p. 2240). If the null hypothesis is true and the ancestral populations of P1, P2, and P3 were equally likely to interbreed without selection differences, then the derived alleles in P3 should match those in P1 and P2 with equal frequency and the D-statistic of the ABBA-BABA test should be zero result. Significant deviations from zero would require an explanation, which could be an introgression from P3 to P1 or P2. Alternatively, from a 'ghost population' PG very similar to P3, which may no longer exist (DURAND et al. 2011, 2040).

Repurposing the ABBA-BABA test to color classes or specific genes: In the study by VICKREY et al. In 2011, the broader ABBA-BABA methodology will be repurposed to address the specific question of whether the domestic pigeon's check pattern was transferred from the Guinea pigeon. In the experimental setup, P1 are barred domestic pigeons, P2 are checkered domestic pigeons. P3 is the exclusively checkered Guinea pigeon. The Wood Pigeon was chosen as the outgroup. P1 and P2 are therefore not different species that mainly reproduce among themselves, but rather different colors of the domestic pigeon. In his studies in 1939, HARMS did not attribute their own racial character to these (p. 11). The calculation requires the identification of transferred (derived) genes or those that are believed to be such. When mated randomly and maintained in symbiosis over long periods of time, the expectation is that 'derived' and putative 'derived' alleles will be distributed equally between barred and checkered individuals.

Significance of the studies for color variations of domestic pigeons

The checkered pigeon in the English Dovecots was given its own status by BLYTH as C. affinis during DARWIN's lifetime, which DARWIN (1868) denied with many arguments. He considered the barred version of the rock pigeon to be the older one. For WHITMAN it was the other way around (p. 49). When repurposing the ABBA-BABA tests to domestic pigeons, checkered pigeons are treated as a separate population. Coming from animal breeding, it's hard to imagine that barred and checkered animals, which have lived in symbiosis for centuries, are systematically different from one another. An exception is the genes that determine the pattern. The regionally delimited Viennese city pigeons, which form a reproductive community, may form a population, but not individual colors from it. Unless there is a strong affinity when choosing a partner for the same pattern. The study cites, among other things, the study of feral domestic pigeons (ferals) by JOHNSTON/JOHNSON 1989 to prove an affinity. This suggests, however, shows that there is a greater preference of bar and check to intermix through their choice of partner. In the fancy, breeding for shows, the standard encourages a pairing of barred and checkered animals. Heterozygous check individuals usually correspond more to the standard expectations with open checks than homozygous ones. This is also an incentive to pair both colors with each other.

On the empirical side: For the gene region in which the patterns are anchored, the D statistics show values ​​close to one. In terms of measurement, these gene areas of the checkered domestic pigeon largely correspond to the gene areas of the Guinea pigeon and check is the central 'derived allele' from the original question. One difference is that no repeats of gene sections (copy number variation) were found in the Guinea pigeon (VICKREY et al. 2011). For the whole genome, D-statistics values ​​close to zero were determined. Positive at 0.021, which is considered an indication of introgression from P3. It is not possible for outsiders to recognize which phenotypes or characteristics are behind the suspected “derived alleles” and how many there are in the sample. In the case of rare genes, drift in the populations will cause problems in clearly identifying “derived” alleles. Based on what is known so far about genetic linkages and correlations and about the pigeon's mating selection, deviations in D values ​​from zero require explanation. As with mutations, they could also be due to chance and sample selection. Rare archaic genes, present or lost in varying proportions across species due to genetic drift, could be confused with derived alleles. Overall, a manageable number of individuals were examined. Significance at low D-values can be achieved with a moderate number of analysed individuals if several gene loci are considered in each case. The mathematical sample size, which is important for the formal significance statement, thus increases multiplicatively.


The proportion of checkered pigeons among domestic pigeons has increased significantly in recent centuries. It is therefore interesting to investigate possible causes and the question of whether the check pattern got into domestic pigeons through mutation during domestication or through hybridization with the Guinea pigeon. From an animal breeding perspective, it is rather questionable whether ABBA-BABA tests can be of any methodological help for this question. Checkered and barred domestic pigeons do not form separate populations. They are different colors of a reproductive community in city pigeons and racing pigeons. Pigeons do not have such a strong affinity when mating within identical colors that, according to previous findings from crosses between breeds and studies of genetic linkages, potentially acquired (derived) alleles remain connected for centuries. Perhaps molecular genetic studies will soon say more about this and/or something different. The question of 'derived' alleles is closely linked to the question of whether one can imagine that mutations repeat themselves. This is assumed to be excluded in DURAND's methodological presentation. If this is the case, then the exclusive existence of the gene in the receiving and releasing population, be it checkered or barred, can be a strong indication, regardless of the value of the D statistic, viewed alone. WHITMAN (p. 19) considered the check pattern to be an ancestral feature of the phylum of pigeons, which was modified in the barred rock pigeon by direct and gradual modifications. If the programming for checks is preserved in the genome, the trait could be activated by parallel selectively triggering mutations, which could explain, for example, a surprisingly rapid parallel fixation of traits in a parallel evolution of separate populations in cichlids (Urban et al. 2020, p. 466). It is possible that parallels can be found in other animal species.


Albin, Eleazar, Natural History of Birds. Illustrated With a Hundred and one Copper Plates, Engraven from the Life, published by the Author and carefully colour’d by his Daugh­ter and Himself, from the Originals, drawn from the live Birds, Vol III London MDCCXXXVIII (1738)

Bechstein, Johann Matthäus, Gemeinnützige Naturgeschichte Deutschlands nach allen drey Reichen, 4. Band Leipzig 1795

Bechstein, Johann Matthäus, Gemeinnützige Naturgeschichte Deutschlands nach allen drey Rei­chen. Ein Handbuch zur deutlichern und vollständigern Selbstbelehrung beson­ders für Forst­männer, Jugendlehrern und Oekonomen, Dritter Band, Mit Kupfern, Zweite vermehrte und ver­besserte Auflage, Leipzig 1807

Brehm, Christian Ludwig, Die Naturgeschichte und Zucht der Tauben, Weimar 1857, Reprint Leipzig 1981.

Carlen E, Munshi-South J. Widespread genetic connectivity of feral pigeons across the Northeastern megacity. Evol Appl. 2021;14:150–162. https://doi. org/10.1111/eva.1297

Darwin, C. R. 1875. The variation of animals and plants under domestication. London, John Murray. 2d edition. Volume 1

Darwin, Charles, The variation of animals and plants under domestication. 2 vols. 2nd edn. New York, D. Ap­pleton & Co. 1883. [first published London, John Murray, 1868]. The writings of Charles Darwin on the web by John van Wyhe.

Dixon, E.S., The Dovecote and the Aviary, London 1851

Durand, Eric et al., Testing for Ancient Admixture between Closely Related Populations, Mol. Biol. Evol. 28(8):2239-2252, 2011

Friedrich II, Das Falkenbuch Kaiser Friedrichs II. Über die Kunst mit Vögeln zu jagen - Süditalien, 1258-1266; After the magnificent manuscript in the Vatican Library. Introduction and commentary by Carl Arnold Willemsen, Harenberg. Dortmund 1980

Goodwin, Derek, Pigeons and Doves of the World, British Museum (Natural History), 2nd edition London 1970

Gray, Annie P., Birds Hybrids, A Check-List with Bibliography, Bucks, England 1958

Haag-Wackernagel, Daniel, Die Taube. Vom heiligen Vogel der Liebesgöttin zur Straßentaube, Basel 1998

Haag-Wackernagel, Daniel, Heeb, Philipp and Leiss, Andreas (2006) 'Phenotype-dependent selection of juvenile urban Feral Pigeons Columba livia, Bird Study, 53: 2, 163 — 170: DOI: 10.1080/00063650609461429; URL: http://dx.doi.org/10.1080/00063650609461429

Kabir, M. Ashraful, Rock-Pigeons in Some Parts of Bangladesh, The Journal of Middle East and North Africa Sciences 2016; 2(3) http://www.jomenas.org

Kinzelbach, Ragnar K. und Jochen Hölzinger (Hrsg.), Markus zum Lamm (1544-1606), Die Vogelbilder aus dem Thesaurum Picturarum, Ulmer: Stuttgart 2000

Kühn, Alfred, Grundriss der Vererbungslehre, 8. Auflage, Heidelberg 1961

McGill Library Archival Collections, Pariser Kröpfer, Peter Paillou (1720-1790)

Santos, C. D. et al., Personality and morphological traits affect pigeon survival from raptor attacks. Sci. Rep. 5, 15490; doi: 10.1038/srep15490 (2015).

Sell, Axel, Critical Issues in Pigeon Breeding. What we know and what we believe to know. Anecdotal, Entertaining, and educational comments on open questions Part 1-7, Achim 2020-2023


Sell, Axel, Pigeon Genetics. Applied Genetics in the Domestic Pigeon, Achim 2012

Sell, Axel, Taubenrassen. Entstehung, Herkunft, Verwandtschaften. Faszination Tauben durch die Jahrhunderte, Achim 2009

Urban, Sabine, et al., Different Sources of Allelic Variation Drove Repeated Color Pattern Divergence in Cichlid Fishes (University of Konstanz), Mol. Biol. Evol. 38(2): 465-477 Advance Access publication September 17, 2020

Vickrey, Anna I. et al., Introgression of regulatory alleles and a missense coding mutation drive plumage pattern diversity in the rock pigeon, eLifesciences.org 2018

Whitman, Charles Otis (1842-1910), Orthogenetic Evolution in Pigeons, Posthumous Works edited by Oscar Riddle, Vol. I, The Carnegie Institution of Washington, Washington 1919. Bird Study (2006) 53, 163–170, © 2006 British Trust for Ornithology

Wikipedia, Columba maculosa. Lip Kee Yap, CC BY-SA 2.0 <https://creativecommons.org/licenses/by-sa/2.0>, via Wikimedia Commons