Pitfalls in traditional and molecular genetic classification of pigeon breeds

Pigeon breeds can be viewed and subdivided from different perspectives.

1. historically, striking characteristics and behaviours are described side by side when depicting domestic pigeons. In Frisch 1763 and in other early writings, these got designations for groups of pigeon breeds such as pouters, tumblers, high-flyers, drummers etc., which still differed from each other in other details within the group. These groupings can still be found today as a classification scheme in the book of standards of the pigeon fancier organisations.

2. the interest of Darwin (1868) in this context was not directed towards the differences of breeds and breed groups, but towards the succession of preliminary stages and preliminary final stages of the development of breeds. This enabled him to illustrate the rapid change of characteristics under the influence of selection.

3. even before Mendel (1865), but increasingly after him, breeds with other characteristics were seen as a possible source for the improvement of breeds or for the creation of new breeds by uniting positive characteristics.

4. molecular genetic classifications of breeds in clusters are made according to genetic similarities identified by DNA analyses. Breeds with particularly high similarity to each other, but with a clear distance to other groups, are grouped together.

Some statements about these studies give the impression that there is little information about the relationships between breeds, especially with pigeons: as unlike with many other domestic species, few reliable records exist about the origins of, and relationships between, each of the breeds” (Pacheco et al., S. 137). As explained below, this underestimates the wealth of information available in the fancier literature. On the other hand, the possibilities of making unambiguous statements are overestimated in view of the breeding activities and multiple crossbreeding between breed groups in the past.


Traditional classifications of breeds according to their utility, appearance and behaviour

In the historical European literature on pigeons, the description of characteristics of breeds and their presumed regional origin are central. So, in Gessner 1555/1557 and in the revised version 1669, in Aldrovandi around 1600, Willughby 1676, Moore 1735 and Frisch 1763. Frisch had already distinguished nine basic types and presented one example for each group pictorially. Field pigeons and Montauben (month pigeons) as ancestors of today's colour pigeons, drummer pigeons, pouters, owls, tumblers, Turkish pigeons as representatives of warty pigeons, Jacobins and fantail pigeons as representatives of the structural pigeons.

Fig. 1: Representatives of two main groups of domestic pigeons by Frisch 1763, Pouters and Turkish Pigeon (Source: Sell: Taubenrassen 2009).

In the German book of standards the groups 'form pigeons’ and 'hen pigeons' have been added. The heterogeneous group 'form pigeons' also includes the large utility pigeons listed as a group in Buffon 1772, among them the giant pigeons already classified as 'Runts' in Willughby 1676 and as Romans in Dürigen 1886. In 2022, the breeding stock of the German BDRG shows 35 pouter breeds from the 'prototype' shown by Frisch, and far more than 100 breeds of tumblers and high-flyers. The classification of the NPA, the American Pigeon Breeders Association, is similar and follows the tradition of other associations.

Historical classifications may be overtaken by development and prove inaccurate. Dixon had already written this in 1851 for some subgroups of tumblers that had degenerated from a flying point of view: they would be called tumblers because they would roll over if they could still fly (p. 118). Associations sometimes react to changes that cannot be overlooked by regrouping. For example, the Coburg Larks and Lynx Pigeons, which were classified as coloured pigeons by Dürigen in 1886, have become form pigeons in the current book of standards, and Strasser as former hen pigeons have also become form pigeons.


Darwin: Changes and emergence of breeds by selection

For Darwin, pigeon breeds were important because he was able to use them as examples to show the rapid change of characteristics through selection. It is about the ancestors of these breeds and about the preforms that still existed at his time. It is not the side by side of breed groups that is of interest, but in this context the differences between the breeds and their predecessors (parent stock). These are indicated in his illustrations in dotted lines. He was able to show the extreme change of characteristics in a relatively short time. In the 'family tree' of the English Short Faced Tumbler, for example, it is shown with a small figure and extremely short beak as the end of a developmental path. Among its direct ancestors are the Common Tumbler, common in England, and, more distantly, the related Lotan Tumbler and the Persian Tumbler.

Fig. 2: English Short Faced Tumbler and its line of development in Darwin 1868 (Source: Sell, Taubenrassen 2009).


The Carrier, with a long beak and relatively high and slender stance, is also at the end of an evolutionary line. Darwin had measured 3.6 cm beak length from the tip to the corner of the mouth in extreme specimens. The Dragon (now Dragoon), the Bussorah Carrier and other Carrier breeds with much shorter beaks are mentioned as predecessors in Darwin’s illustration (cf. Sell, 2009, p. 124).

Fig. 3: English Carrier in Darwin 1868, and development of beak length over time (Source: Sell, Taubenrassen 2009).

Whether the development of the beak length in both breeds was due to selection alone remains to be seen. For in the historical literature there are indications and evidence that the present French Bagdette, which existed under other names at that time, was involved in the development of the English Show Carrier, and, in the case of the beak length of the English Short Faced Tumbler, also owls.


Aldrovandi in Europe did not know the fantail in 1600. In 1669 it appears in the version of Gessners revised by Horst as Cprian peacock tail. It was fetched from Holland by local noble lovers for a lot of money. The number of tail feathers is given as 26 instead of the 12 of a normal domestic pigeon. This is also the number quoted by Darwin with reference to Willughby (1676). Moore (1735) names 38, Buffon (1772) 32 and Boitard and Corbié (1824) 42. In England, according to Darwin, it was not so much a question of the number, but also of the general posture (general carriage). Today, this has changed even further with extremely short backs (Fig. 4).


Willughby (1676) describes them with at least 26 tail feathers instead of the normal 12, Moore (1735) with 38 and Buffon (1772) with 32. Darwin quotes Boitard and Corbiè (1824) who counted 42. In England, however, it was not so much about the number, but also about the general posture (general carriage). This has changed even further today with extremely short backs according to Darwin (Fig. 4, 10).


Fig. 4: Fantail at Darwin and development from Buffon (1772) via P.J. SELBY (1835/1843) to today on the cover of the book Taubenrassen.


Creation of new breeds and transmission of traits across breed groups

The fact that traits - positive and negative - are hereditary was a firm knowledge of practical animal breeding even before Mendel. Under the keyword "Livestock Breeding" in the Book of Inventions, Leipzig and Berlin 1864, we find the following statements:

"In relation to performance ... the individual species, horse, goat, cattle, etc., are not only, but also individual branches of these species among themselves again very different. The name breed means as much as a variety distinguished by special characteristics common to all members of the breed. We only find breeds among those animals which man has taken into his immediate environment, made into domestic animals and accustomed to his service"... "But it is just as certain ... that man has it in his power to form new breeds according to his will"... (in German language, p. 204ff.). And further on breeding strategies: "The animal breeder can now preserve the existing breeds or change them in a refining way. A breed is preserved by mating the best animals within it; it is improved by inheriting forms or characteristics which it lacks for a desired purpose or in general, namely by mating animals of different breeds in which those desired characteristics appear sporadically" (Buch der Erfindungen 1864, p. 204, quoted at Sell, Taubenrassen, p. 191).

For many pigeon breeds this is documented in detail by Boitard and Corbié already in 1824. In Germany, originally inconspicuous drumming pigeons were adapted to the taste of the breeders by crossing with colour pigeons. Thus, the newly developed drumming pigeon breeds were long referred to in the literature as 'bastard drumming pigeons' (Neumeister 1837 and Neumeister / Prütz 1876).



Fig. 5: Drumming pigeons and bastard drumming pigeons in Neumeister 1837, Neumeister /Prütz 1876. Sketch of the interrelationships of breed groups (Source: Sell, Taubenrassen 2009).


The group of tumblers and high fliers is particularly receptive to traits originally associated with other breed groups. High stance, long necks and long beaks due to influence of French Bagdettes (Fig. 6), short and thick beaks due to Owls, feather ornaments of Drummers and Fantails, colourings of Colour Pigeons.

Fig. 6: German Magpie Tumblers in Schachzabel 1910, Magpie Tumbler in Lyell 1887 and English Magpie Tumbler from 1938, illustrated in Levi 1969 (Source: Sell, Pigeon Genetics 2012).

In Magpie Tumblers, pigeons modified by crossbreeding first appeared as a 'modern type of the breed' before they became accepted as successors to the old breed. This was also the case with Maltese Pigeons, Kassel Tumblers and others (Sell, 2009).

Fig. 7: Maltese of old and ‘modern type’ at Dürigen 1868 and 1906, source: Sell, Taubenrassen 2009.


Strasser and Lynx pigeons have also not developed from the pigeons shown in Lavalle und Lietze 1905 by selection in figure and size alone.

Fig. 8: Change of the Lynx Pigeon from a Colour Pigeon to a Form Pigeon and of the Strasser from a Hen Pigeon to a Form Pigeon (Source: Sell, Pigeons Genetics. Applied Genetics in the Domestic Pigeon, 2012).


A source of breed mixing is also the transfer of newly discovered colour factors to other breeds and breed groups. Discovered in 1945 in roller pigeons, for example, the 'Reduced' factor has been rapidly transferred in the USA to Genuine Homer, Giant Homer, Racing Homers, Long Faced Tumblers and other breeds. It came to Europe via such Long-Faced Tumbler crosses. In the Dortmund area the breeding partnership Christoph Mooren and Thomas Schmidtmann created reduced Cologne Tumblers from these. From these the hereditary factor was transferred by Norbert Dietrich to German Modenesers and by Fritz Muchow to Colour Pigeons in Thuringian Shield Pigeons. Thomas Voss transferred the factor to European Racing Homers. In the meantime, the factor is present in Owls, Drumming Pigeons and other breed groups.

Fig. 9: Transfer of the Reduced hereditary factor from roller pigeons in the USA to many breeds within a few years (Source: Sell, Critical Issues in Pigeon Breeding, Part IV 2021).


The transfer of other factors can be traced with similar accuracy in the hobby literature. For example, for the factor indigo uncovered in crosses in the USA in the 1940s, which was later shown to be found in high performing European racing pigeon strains as well (Sell 2012, 2019).

Fig. 10: Transfer of the hereditary factor indigo in a few generations to other breed groups such as high-flyers from Racing Homers in the author’ loft and Fantails (Source: Sell Pigeon Genetics. Applied Genetics in the Domestic Pigeon, 2012).

At the same time as the colour factors are transferred from one breed to the other, genes from other gene regions are also found in this and other colour classes through crosses with them.


Molecular genetic classifications

Forms of representation

Following a study by Stringham et al. (2012), a number of other studies have appeared on pigeon breed similarity and classification in clusters and representations in dendrograms. The study used 32 unlinked microsatellite markers to genetically characterise 361 individuals from 70 domestic pigeon breeds and two free-living populations. Similarly extensive, and partly building on it, the study by Pacheco et al. (2020). Confidence-building that the tumblers and high-flyers shown here in the excerpt from the dendrogram given by Stringham, such as Birmingham Roller (BIR), Tippler (TIP), West of England Tumbler (WOE), remain largely together, which confirms or seems to confirm the assumed commonality of origin.

Fig. 11: Example of a dendrogram. Excerpt from Stringham and others (2012). A breed classification from the English Cropper (ENP) to the American Show Racer (ASR).

Similarity by common ancestry or recent crosses?

A proven similarity does not mean a reliable statement about the origins of the breed. Similarity is determined between current stock. The similarity may be due to common ancestors, it may also be due to interbreeding between the breeds studied or between preforms of these breeds.  It is therefore not surprising that in a study presented by a Polish research group (Biala et al. 2015), a major finding was that the breeds studied were subject to permanent genetic mixing.

Lexicographic classification versus classification by measuring similarity at multiple gene loci.

The newly formed clusters are no substitute for grouping in pigeon fancy. The English Cropper (ENP) will not be grouped with the Stargard Shaker (ZIT) in the fancy, despite its grouping in the dendrogram (Fig. 11). Historically, a single characteristic decides on the grouping of pouters, drumming pigeons and tumblers/highflyers. Pouters blow up their crop, drummer have a special vocalisation. They are still pouters or drummer respectively, even if they differ in size, colouration, posture and feather ornaments (cap, foot feathering, etc.). Even with the formerly classified giant pigeons, only this one criterion decided with the size.

The molecular genetic clustering is different. Here, a multi-dimensional approach is chosen to define a cluster by determining the gene regions to be included in the analysis. In Frisch, genes responsible for drumming, the inflation of the crop or the tendency and ability to fly high and tumble were the only ones responsible for group membership. In molecular genetic clustering, this is in each case only one trait among many, which is only indirectly taken into account, if at all.

The importance of the individuals selected for analysis

For the results of the analyses, it is important which individuals represent the respective races. Dominant hereditary factors, for example, can be transferred from one breed to another within a few generations. In the case of recessive ones, it takes somewhat longer. Even experts of the breed are often unable to detect differences from the original breed a few generations after crossbreeding. Individuals of one breed can therefore differ considerably in the measured genetic distance, which is also pointed out by Pacheco et al. in the Supplement for their study. Breeders from Holland presented some years ago black-winged copper Gimpel Pigeons (Archangel) into which King Pigeons had been crossed a few generations earlier to obtain more body mass. Even experts could not find any differences to the pure breed. Some breeders will consider the first cross-breeding results of a mating of a peak-crested golden-white winged cock of the Gimpel Pigeons with a black tigered short-beaked highflyer hen as differently coloured Gimpel Pigeons. Some of the plain headed and peak-cested golden white wings originating from the first back-mating upon the golden-white-wing cock were difficult to distinguish from pure-bred birds by type, beak length and colouration, even by experts (Fig. 12).






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Fig. 12: Mating of a peak-crested golden-white-wing Gimpel cock with a plain-headed black-tigered highflyer hen with two of their young. On the right, a female of the first back-mating to the golden-white-wing cock at the author's loft (source: Sell, Genetik der Taubenfärbungen, Achim 2015).

If at first only the genome of birds coming directly from crosses has changed, these changes will affect the whole breed through mating with other flocks over the years. Reasons for such cross-breeding can be colouration, feather ornaments, flight ability or vitality in general (Sell, 2012, 2019). Especially in rare breeds with frequent breeding borrowings from other breeds, a molecular genetic finding therefore only represents a snapshot.

Irritations in the representation of dendrograms

There are various algorithms that lead to different dendrograms and thus give the viewer different impressions of the network of relationships between the breeds. Some irritating assignments to clusters are due to the reasons mentioned above, as Pacheco et al. also noted with reference to background information on the animals studied. Assignments of breeds with large genetic distances to each other point to still other methodological reasons. So do the cases where it could be assumed from the small genetic distance that the breeds would quickly find themselves in a common cluster. When this does not happen, it may be due to third party relationships. The environment of the objects also influences the assignment. Deletions, or even additions of races, can trigger a complete reorganisation of the dendrogram.

Concluding remarks

For Darwin, the historical perspective of races and the appearance of preforms (parent stocks) were of particular importance for showing rapid changes through selection. He was able to show this exemplarily with outwardly extremely different breeds.

In pigeon fancying, the differentiation of breed groups was and still is about a brief characterisation, which was usually based on a single, easily identifiable characteristic that was considered significant. As early as Frisch 1763, these were and still are pouting, drumming, tumbling, warts, special feather structures, etc. Genetic similarities in other traits are considered secondary.

One justification for classifying breeds via molecular genetic studies was that in pigeons, unlike other domestic animal species, there are few reports on origin and relationships between breeds (Pacheco et al., p. 137) are not true. The study itself refutes this by reference to relevant literature and this is shown by the exemplary references above. Without knowledge of the literature, it is easy to misinterpret fresh traces from recent interbreeding as old relationships. Conversely, old ones will be overlooked because they have been overwritten by more recent crosses. The molecular genetic analyses obtained are each a snapshot in time. The constant over time for some breeds is only the breed names. Nevertheless, the results on genetic differences of breeds remain interesting. In the medium term, the changes in breeds over time could be monitored. And this could possibly be done by including historical specimens.


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