This article was published in Flare (Journal of the IBC) - November/December 2009, Volume 43, No. 3.

For the past several years, both authors have extensively worked on metallics in their breeding programs. Their experiences and thoughts on the metallic trait were written down in several articles ("Copper gold" (2003) [1]; "Metallics and Masks" (2005) [2]; "Understanding metallic genetics" (2006) [3]). Both authors decided to team up in order to share their thoughts and experiences on "dragons".

It is clear that the development of the metallic betta was the catalyst of a true "bling-bling"-fever among betta hobbyists worldwide. Metallics have greatly influenced the betta hobby and opened doors for many breeders to develop new color varieties and combinations. Following the metallic betta which made such a prominent mark on the betta hobby, a new variant came hard on its heels. The unique appearance of this so-called "dragon" betta is characterized by a thick, solid silvery/white metallic layer which almost resembles armor. "Dragons" captured the imaginations of betta breeders the world over, leading to the hype that has been humorously termed "dragon fever". But what exactly are "dragons", where did they come from and are they really different from our metallics?

Black "dragon" [7]

The origin of the "dragon" betta

According to Pichet (Interfish breeder team - Thailand), the first "dragons" were created by using a "super" red plakat, a red copper plakat and Betta sp. mahachai [8]. With some difficulty, the wild Betta sp. mahachai was crossed to the red copper plakat with the intent of preserving the Betta sp. mahachai traits in the offspring. Then a young male from this spawn was crossed to a super red plakat female. The third step was a mother x son backcross: the super red female was crossed to one of her offspring in a method commonly referred to as 'line breeding'. Although the body shape and finnage of the fry was not very good and some of them even showed malformations in the body, the color was there and the first dragon pair was born! The first "dragons" were developed by Mr. Tea. The "Red dragon V1" were first presented to the Thai public in the December 2004 edition of a magazine called "Fancy Fish" by the Interfish breeder team [9].

The shiny iridescent layer of the red "dragon" V1 was not completely covering the body and it was Mr. Somchat (Interfish breeder team - Thailand) who used these fish to improve the color and finnage into the red "dragon" V2. In 2005, the first dragons were shown at Aquarama by the Interfish breeder team who also first introduced the red "dragon" to the rest of the world. At this stage the fish was still shortfinned and although it did not take home any prizes, many Asian breeders were impressed by its appearance.

The "Armadillo" strain

Around the time the Interfish breeder team was amazing the world with their new "dragon" bettas, a strain called the "Armadillo" was being produced in the United States by Victoria Parnell-Stark of Bettysplendens.com. The initial idea for them came from a book by Walt Maurus [10], in which he included a photograph of a very unique looking betta that was the result of crossing Betta splendens with wild bettas, such as Betta imbellis. This beauty was called the "Neon" betta. Determined to reproduce the anomaly, Victoria Parnell-Stark acquired several pairs of wilds from Thailand in a shipment that included Betta imbellis as well as Betta sp. mahachai. As with the results already described above, the Betta sp. mahachai crosses proved not only difficult but temperamental, and it was only with some trial and error that a successful spawn was recorded between a Betta sp. mahachai and a green Betta splendens. However, the resulting fry were never particularly hardy or prolific.
There was considerably more success with the Betta splendens x Betta imbellis crossings, and some very interesting, highly iridescent and masked fish were beginning to appear. The first generation yielded many green/red and multicolored bettas, some with highly iridescent scales and brightly colored gills. Sequential breedings enhanced the effect, so that within a few generations bettas with heavier iridescence and fully masked faces were beginning to appear. The Betta sp. mahachai x Betta splendens hybrids were not faring so well, so before their influence was lost entirely they were absorbed into the line of Betta splendens x Betta imbellis hybrids. In 2006 the first "Armadillo" bettas were offered for sale, and it was not very long before betta enthusiasts were noticing similarities between the "Armadillos" and the "dragons". Indeed, their origins followed a very correlative formula, with the differences (particularly in color) probably being attributed to the amount of Imbellis influence in the "Armadillo".

"Dragons" - Back to the roots!

Betta imbellis, Betta smaragdina and Betta sp. mahachai naturally have a higher degree of iridescence on the scales compared to Betta splendens which probably is an adaption to the murkier water in their natural habitat. Both "dragons" and "armadillos" described above are characterized by a thick metallic iridescent scaling which almost resembles body armor. When we have a look at the origin of metallics and "dragons" we see quite a lot of overlap. Outcrossing domesticated bettas to Betta imbellis gave rise to the metallics which in turn also have an important contribution to the development of the "dragon". This shared origin mostly likely explains the similarity between metallics and "dragons" but in case of the latter the additional influence of Betta sp. mahachai seems to be the key to the thick shiny armor-like metallic scaling. When we compare wildtype Betta imbellis and Betta sp. mahachai we can already see the iridescence is shinier and thicker in the latter.

 When we take these four (sub)species theoretically the following crosses could be made:
·   Betta splendens x Betta imbellis
·   Betta splendens x Betta smaragdina
·   Betta splendens x Betta sp. mahachai
·   Betta imbellis x Betta smaragdina
·   Betta imbellis x Betta sp. mahachai
·   Betta smaragdina x Betta sp. mahachai

With exception of Betta splendens x Betta imbellis crosses, not much is documented about whether all these species really can interbreed without problems. When we look at the origin of the "dragon" as described above, accoring to Pichet it was not easy to cross the red copper betta x Betta sp. mahachai. Please note that copper is a metallic phenotype which was created by crossing Betta splendens to Betta imbellis, however how many generations this particular fish is seperated from this initial hybridization is unknown. Strikingly, Victoria Parnell-Stark also reported difficulties with the Betta splendens x Betta sp. mahachai hybrids. In both cases it seems that a mixture of these three species was necessary in order to create and fixate a healthy, viable line. So why where some of these crosses difficult? Was this is just a coincidence or could it be that certain species were evolutionary more separated from each other? Does the similarity in appearance between Betta imbellis and Betta sp. mahachai and the readiness to interbreed mean that they are evolutionary closer related to each other than to Betta smaragdina and/or Betta sp. mahachai? Based on their phenotype Betta smaragdina and Betta sp. mahachai also show quite some similarities, does this implicate that they are more easily easily intercrossed than to Betta splendens and/or Betta imbellis?

Some other unanswered questions/remarks to keep in mind with respect to this subject are:
(1) Does it matter which sex we use to interbreed species? Are the results different when we breed a male of species A with a female of species B or the other way around?
(2) Are F1 hybrids of the above mentioned crosses normally fertile, and is this sex dependent? It is known that F1 hybrids of two species which are further apart in the phylogenetic tree could be infertile. Does this account for both sexes or is it in some cases possible to use hybrid males or females to cross back to one of the species used in the initial cross?
(3) Can hybridism function as a genetical "bridge" between two distinct species? The idea of the later hypothesis would be that the F1 hybrid offspring of two different species might function as a genetical "bridge" for the interbreeding of two species which are evolutionary further apart.

"Dragons" -  Genetics?

We know that the first metallic bettas were created around the year 2000. It took about 5 years before Dr. Leo Buss provided published several papers which provided us with more information and insight about the genetic make-up of this phenotype [15-18]. By now the same time-span has passed since the first "dragons" were developed but so far not much is known about genetic make-up and inheritance of this phenotype and thereby leaving us with quite some unanswered questions:

·   What  genes determine the "dragon" phenotype?

We can clearly see that the unique appearance of the "dragon" clearly affects the iridescent layer. So far several genes have been characterized which affect the iridescent layer, for example the classical iridescent colors steel blue (blbl), turquoise (BlBl) and royal blue (Blbl), metallic or yellow reflecting iridophore (the wildtype variant of the reduced yellow iridophore locus) [18] and the spread iridescence gene (Si) which is responsible for the increase in density and distribution of the iridescent color.
In 2007, Joep van Esch of Bettaterritory.nl made some experimental crosses with "dragons" in his fishroom in order to learn more about the heredity behaviour of this trait (see pedigree below). After crossing a "dragon" to a regular metallic (BT171007A) he observed that all offspring had a copper phenotype. Considering the fact that  a copper phenotype (homozygous metallic steel blue, blbl ++) can only be obtained when the offspring obtains one steel blue (bl) and one metallic (+) allele from each parent [3], this result suggests that both traits may play an important role in the unique appearance of the "dragon" phenotype. Interestingly, the metallic scaling of the offspring seemed more thick and solid than observed in regular metallic fish. This characteristic was also passed on to the offspring when a heterozygous “dragon” was crossed to a heterozygous metallic fish (BT271107). The offspring resulting from crossing two heterozygous metallic sibblings (BT171007B) resulted in homozygous, heterozygous and non-metallic offspring without this unique appearance which was observed before. Off course we have to keep in mind that these findings were purely based on macroscopic observations (with the naked eye). Nevertheless, these crosses show that the “dragon” trait is hereditable and suggests that it behaves quite dominant as it already clearly affects the iridescent layer in heterozygous fish.

In the breeding reports of Pichet and Victoria Parnell-Stark outcrossing a metallic fish to Betta sp. mahachai seems to have been the key to the thick shiny armor-like metallic scaling of "dragons". Dr. Leo Buss has reported that domesticated metallic bettas and wildtype Betta imbellis possess yellow reflecting iridophores [16] but whether this also hold true for Betta smaragdina and Betta sp. mahachai to our knowledge is unknown. It is known that it is possible that several variants of a certain gene (polymorphisms) can exist within a population of a certain species. As Betta splendens, Betta imbellis, Betta smaragdina and Betta sp. mahachai all share a common ancestor, there is a chance such gene variants also exist among these (sub)species and that they were combined by crossing the different (sub)species. Which gene(s) and/or gene variant(s) exactly are responsible for the unique appearance of "dragons" is difficult to answer without any microscopic examination of the chromatophores themselves and a determined breeding program.

·   Do blue "dragons" exist?

The characteristic thick, silvery/white metallic layer on the body is typically found red, yellow, orange, white and black "dragons". In case of red, yellow and orange "dragons" this layer can be observed in both light- and darkbodied fish.
Interestingly, so far this characteristic silvery/white metallic layer has not been observed in darkbodied bettas with a classical iridescent color (steel blue, turquoise or royal blue) of the body and finnage. The picture shows an imaginary example of what the authors mean with this.
In case of a dark bodied "dragons" with a classical iridescent base, the silvery/white appearance seems to be masked although based on macroscopic observation (with the naked eye) their scaling definitely seems to differ from a regular metallic fish by being more solid. There have been reports of red and black dragons with a blueish shine on their scaling but so far no real blue "dragons" with this typical thick, silvery/white scaling have been developed. It would be interesting to find out whether it is possible to obtain a the thick, silvery/white metallic layer back after crossing a blue or copper "dragon" as depicted above to a red, yellow, orange or black betta.

·   Is it possible to breed non-masked "dragons"?

In general the "dragon" phenotype seems to be associated with the iridescent layer covering the head, the so-called mask trait. It is known that the metallic and mask trait are caused by seperate genes and can work independently. With the exception of a few marble "dragons", so far to our knowledge no non-masked "dragons" have been developed but off course this may change in the future.

To conclude.......

Although the "dragon" phenotype has been introduced into our hobby for about 5 years ago, its possibilities still seem unlimited and new color combinations and patterns are continuously being developed. But there still are a lot of questions to be answered. It still is not clear how we should define the true "dragon". In order to answer this much more research has to be done. A detailed microscopic examination of the chromatophores, as previously done with the yellow reflecting iridophore [15-18], in combination with a determined breeding program might provide more information with respect to which gene(s) and/or gene variant(s) are responsible for the unique appearance and inheritance of the "dragon" phenotype.

Welcome in a new era in the world of "bling-bling" bettas!!
 

Acknowledgements:

We would like to thank Dr. Leo Buss (USA), Stefan Psarakos (Australia) and Ursula Bosnjak (Austria) for their suggestions and critical review of this article.

References/credits:

1. Parnell-Stark, V., "Copper gold”, 2003.
2. Parnell-Stark, V., "Metallics and Masks”, 2005.
3. van Esch, J.H.M., "Understanding metallic genetics”, 2006 and published in FLARE magazine, Vol. 41, No. 6., 2008.
4. Bred by Tada palakul (Smilebetta) - Thailand.
5. Bred by Sanya Ponpal (Interbettas) - Thailand.
6. Bred by Rung Keereelang (Banleangbettas), owned by Robi Iskandar (Robi2000) - Singapore.
7. Bred by Robi Iskandar (Robi2000) - Singapore.
8. Pichet (Interfish Breeder Team - Thailand), personal communication through email, 2008.
9. Fancy Fish Magazine, Thailand, Vol. 5, No. 50, 2008.
10. Maurus, W., “Bettas a complete introduction”, ISBN 0-86622-288-x, 1981.
11. Parnell-Stark, V. (USA) www.bettysplendens.com.
12. Panitvong, N., International Betta Congress (IBC) - Species Maintainance Program (SMP).
13. PIBK2005, Malaysia, 2006.
14. Bred by Vu Minh Chuong - Vietnam; www.bettasaigon.com.
15. Buss, Leo W., "Structural Color", Bettas&More - FAMA, March 2005.
16. Buss, Leo W., "A 'New' Iridophore Color", Bettas&More - FAMA, September 2005.
17. Buss, Leo W., "Inheritance of Metallic Trait", Bettas&More - FAMA, November 2005.
18. Buss, Leo W., "Naming the new metallic gene", Bettas&More - FAMA, March 2006.
19. Picture adapted by Joep van Esch, based on a non-metallic royal HM PK from BT161209.
20. Bred by Le Anh Tuan - Vietnam; www.bettasaigon.com.
21. Bred by Kiat Inthamu (Muangkhonbettas) - Thailand