A Discussion of the Micariff Phenotype; misconceptions and facts

MONDAY, MARCH 30, 2015

A Discussion of the Micariff Phenotype; misconceptions and facts

© Alan S. Bias Permission granted for nonprofit reproduction or duplication of photos and text with proper credit for learning purposes only. March 30, 2015 March 31, 2015 (Updated)

Blond Micariff, courtesy Micke Norberg

Grey Micariff, courtesy Micke Norberg

According to California breeder and IFGA member Mike Khalid, the Micariff phenotype was created in Sri Lanka by two breeders from his homeland, Michael Cole and Denis Ariff. The name itself a truncation of Michael's given name and Denis' surname: MicAriff. While Micariff is most often associated with a reduced melanophore blond (b) phenotype, it is also recognizable in wild-type grey with increased melanophores. As a pedigree stock breeder I am hesitant to assign specific genes to produce expected result (phenotypes). If for no other reason modern research and breeding’s have shown the many phenotypes are not the product of “single genes”, rather the product of “multiple genes” or mutations upon them in combination or linked complexes. To this reasoning, as in all my writings, will refer to a gene(s) expression as a trait.

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The required traits needed to produce the Micariff phenotype have long been the subject of conjecture based on both purebred and outcrosses maintained by breeders. Results from each suffering from lack of scientific method needed over multiple generations to provide proper ratio’s for analysis. Yet these breeding’s can still tell us much from visual records. The two most prevalent theories having arisen in Europe and Asia where Micariff’s have a stronger following by serious breeders in comparison to North America. The Asian version while lessor defined basically states that the end result is the product of a metal trait, likely Metal Gold (Mg) + Snakeskin Body (Ssb) + Snakeskin Tail (Sst). [Note: Hereafter Ssb / Sstwill be referenced singularly as Ssb]. The better defined European version asserts a product of homozygous Stoerzbach1 (s) + homozygous snakeskin (Ssb): Males = XSsb YSsb ss mgmg and females = XSsb XSsb ss mgmg. It should be taken into account here that many European breeders do not make a genetic distinction between Stoerzbach and several other metal traits as is common in Asia and North America.

Grey Micariff female, courtesy Myster Green

Blond Micariff female, courtesy Raj Seshagiri

Hypothesis: In response to both of these theories, I state each as being only partially correct. I propose: That Metal Gold (Mg), Stoerzbach (s), Snakeskin Body (Ssb) all in homozygous state, and a “missing trait” are required to produce the original metallic Micariff phenotype in full body expression. That this missing trait is required in conjunction with Steorzbach a "full body" trait acting as a precursor to effect the "metallic gold Micariff Page 2 of 21

phenotype" (hereafter metallic) in both body and finnage. As a result two actual expressions are commonly produced when Stoerzbach is lacking or in heterozygous form; metallic and non-metallic body (hereafter nonmetallic). [Note: It should be stated that in a true breeding metallic strain females under this hypothesis would need a genotypeXSsb XSsb ss mgmg in females and XSsb YSsb ss mgmg in males. For for producing a percentage of metallic a genotype heterozgous for stoerzbach in either males or females would suffice. For production of non-metallic a genotype lacking Sterozbach in one sex or heterozgous for stoerzbach in both sexes would suffice.]

Ssb

Homozygous metallic; X

Ssb

Y

ss mgmg, courtesy Mitchell Weintraub

Ssb

Heterozygous metallic offspring; X Y

Ssb

Heterozygous non-metallic; X

Ss mgmg, courtesy Mitchell Weintraub

Ssb

Y

Ss mgmg, courtesy P. Shaddock

Discussion: When discussing genetic makeup and expression of color & pattern in guppies, one should undertake an understanding of what we are actually seeing with the naked eye. You quickly come to the realization we are not looking at a colored fish, but rather it's reflective qualities. The Bagnara Dermal Chromatophore Unit was first published in 1968. For nearly four decades it was for the most part ignored by both the scientific community and that of the Domestic Guppy breeder. Though the latter Page 3 of 21

often stumbled upon it in breeding's and produced visible results without understanding the genetic implications. Longtime successful stock breeders breed by eye; through the power of observation to achieve if not understand results. We are visual thinkers to varying degrees. Little different from a musician who plays by ear. This shared trait among breeders has met stiff resistance in acceptance by many. Yet even the scientific community has slowly started to support what we as breeders have known for centuries; that successful breeders have an inherent and often above average ability to observe the natural world and create domestic results (Arnheim 1969 & Grasseni 2004 & 2005). Our shortfall in the eyes of science has been an inability to communicate this understanding to others, both outside and among our peers. In the following simple visual aid, used in support of Bagnara's work on color study, we have the basis of his theory as published in multiple papers over a period of two decades. [Note: color added to black and white photo to reflect chromatophores.]

The Dermal Chromatophore Unit; Bagnara (1968) & Bagnara et. al. (1973), reprint from Grether (2004)

[Excerpt Grether (2004): The dermal chromatophore unit... ...of a hypothetical green frog, showing how the xanthophore, iridophore, melanophore and underlying fascia layers interact to determine the overall colour of the animal. Wavy lines depict the paths of light of differing wavelengths through the cell layers...] An understanding of the Dermal Chromatophore Unit in conjunction with an understanding of genetic associations between melanophores - color pigments - iridophores has much to offer serious breeders. The two will help plan your mating's and alleviate trial and error through random breeding's. In the end help understand your results, both positive and negative. Even though your goals or that of a show standard may call for a specified result, is it in reality genetically possible? Tank space is too precious to "recreate the wheel" in reproducing the failures of our predecessors. Yellow color pigment is often very motile and hard to define by the naked eye unless put under a microscope to determine composition. From a breeders perspective I tend to make a couple basic assumptions in associating yellow traits when it comes to yellow color pigment cells. Page 4 of 21

The assumption is based on perceived mode of inheritance and motility. If autosomal dominant or recessive, likely metal gold. If sex-linked, likely yellow color pigment. Even this is not guaranteed. There are so few identified “yellow color pigment” traits in the body. If there is an “expected norm” it might be: 1. Yellow color cells are often very small and round. 2. Red color cells are larger and less defined. 3. Wild-type orange seem to fall in between in both size and shape. Working from memory, wild-type yellow cells are most often punctate; small & round. They can also be dendritic; large with arm-like extensions. The latter can result in regions with dark dense and motile yellow coloration. Population type (punctate or dendritic) and numbers often differ depending upon a lower layer of leucophores or iridophores in both body and finnage. When high concentrations of either leuchophores or iridophores appear absent to the naked eye (clear finnage) dendritic & punctate are often found together close to the body. As you extend further out on finnage dendritic reduce in numbers, while punctate continue. Even then, this is subject to both type and concentration of melanophores. Both red / yellow cell placement and population is in interaction with melanophore types and location. Both punctate and dendritic yellow color cells are found in various locations in body and finnage. Punctate throughout the body and dendritic concentrated in specific locations. Based on breeding results, trait definitions, and an understanding of trait composition it seems dendritic are the most motile. Punctate are scattered about having less direct interaction with melanophores. Dendritic tend to collect around melanophores. The primary difference often determined by leucophores (white), iridophores (silver / blue) or neither underlay color pigment cells. [Note: Motility of yellow color cell pigments over iridophores or leucophores allows for increased expression by positive selection.] The two best examples for understanding yellow color cell population and color type are Schimmelpennig Platinum and Full Platinum strains. Schim Plat is a “fixed” trait, likely in complex. Yellow coloration in Schim Plat is comprised of evenly dispersed punctate cells and areas of collected dendritic cells. Line of decent plays a large role in Schim Plat composition. Schim Plat is a full body trait derived from Vienna Emerald Green. Vienna Emerald Green in turn derived from “wild-type”. Wild-type is comprised of both silver / blue iridophores and Le in well-defined zones of regulation. While the Schim Plat has distinct zones of Le regulation, it is overall a silver / blue iridophore full body trait. [Note: Schim Plat in outcross often expresses reduced levels of Mg yellow coloration in F1 offspring. This in itself is suggestive that we should view Mg not as a “yellow / gold iridophore”, but rather as a “yellow / gold color pigment trait”. There is limited evidence pointing to the existence of either red or yellow iridophores.] Full Plat (NiII + Le) is the product of combination, easily segregated in outcross. The increased expression of Le results from enhancement through combination with a full body trait; NiII. Yellow coloration iin much of the body is predominantly comprised of evenly dispersed punctate cells over white. Iridophore regulation, while present, is minimal. In IFGA circles the Micariff does not command the respect of many breeders as having what is needed to produce a fish worthy of competition in show. Many breeders have attempted to do so, with marginal results, Page 5 of 21

and in the end abandoned the process in favor of more reliable traits to produce a Yellow Guppy. Others have incorporated selected traits into existing strains with positive result. Melanophore reduction will result in increased visibility of color pigmentation; i.e. Blond Yellow or Blond Full Gold appear more yellow than grey counterparts. However, without a corresponding increase in reflective qualities, color may become flatter than grey counterparts. The blond Micariff Guppy is a phenotype that is either loved or hated by individual breeders. It has contained in its genotype in homozygous form the traits needed to produce a Yellow Guppy. Although this is often with corresponding reduction in overall finnage. An outcross can result in near complete loss of yellow color pigment or Mg if you lack an understanding of the genetic composition. The traits most often favored in production of yellow strains involve combinations of metallic iridophores; Metal Gold (Mg) or Schimmelpennig Platinum (Sc), melanophore reduction or masking; Blond (b) or Albino (a), metal enhancement; Stoerzbach (s) or Full Gold (NiII + Mg), and red color pigment removal; Asian Blau (Ab). Due to the very nature of yellow color pigment interaction with melanophores and iridophores, results while impressive have fallen short of achieving a full Yellow Guppy. By some considered genetically impossible and likely so without the further identification of traits in the future. While Stoerzbach and Snakeskin are both considered full body traits, it is long known by breeders that neither the combinations of ss + Ssb or Mg + Ssb will in themselves produce a “true” Micariff phenotype. At best each will produce a “co-expression” of either combination not effecting a full body metallic Micariff expression. Numerous breeders are aware that F1 offspring from either sex Micariff parents in reciprocal outcross will produce a snakeskin phenotype. This in itself is suggestive of Ssb in either sex being in a homozygous state. In turn, sib-breeding of F1 stocks will again recombine in subsequent generations to produce Micariff’s with increasing numbers. Here in outcross and re-combination of s and Ssb is where we first see clear evidence of our missing trait. In various Micariff phenotypes, you will notice distinct variations in color and pattern. Some are more metallic in nature, while others have higher expression of "flatter" colored leucophores. It is common for autosomal traits to produce partial expression in heterozygous fashion. With high expression of leucophores vs. iridophores a red shoulder stripe is often apparent, especially in blond males. This is often referred to as an "Old Fashioned" shoulder stripe; a by-product of Vienna Emerald influence. When faint lavender colored the result of Purple Body Mutation (Pb). When red, the result of X-link red color pigment. In all cases, there is a direct correlation between increased anterior iridophores and a reduction or complete masking of the shoulder stripe.

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Outcross has led to the further development of several other Micariff phenotypes over the last couple decades by both Domestic Guppy Breeders and Commercial Farms. Under the strain names of; Berlin Buttercups, California Golds, German Yellows, Tequila Sunrise, Sunset Guppy, etc… Each the result of various states of zygosity involving the original foundation traits &/or addition of new traits to include; Half Black (NiII), Xlink red color pigment in finnage, Lazuli (Lz), and Japan Blue (Ca)2. Sunset’s are the simple product of the inclusion of X-link red color pigment in heterozygous or homozygous fashion, as can be demonstrated by breeding results on a Punnett square. They are not the product of a distinct “Sunset Micariff” allele as has been proposed & promoted by some breeders. Red color pigment was initially infused as a heterozygous X-linked gene. As result of crossover red color pigment can, and often of does, become Y-link. Expression in Sunset’s will vary between heterozygous and homozygous state. Where is our evidence of a full body trait and zygosity impacting expression of red & yellow color pigment in a Sunset Micariff? It lies in the pattern expressed by the two chromatophores; a clear delineation in coexpression of the red (erythrophores) and yellow xanthophores). In most instances yellow color pigment in guppies, depending upon linkage and zygosity, is epistatic to red. The end product being either yellow or orange (wild-type orange). In the Sunset Micariff, while there may be minimal blending of the two color pigments, the expected result is clear delineation; a repulsion of red away from yellow to the exterior edges of finnage. To produce such a result, a “full body” trait is required. In example: NiII produces a similar repulsion between yellow color pigment and black melanophores in finnage. Thus, demonstrating its effect as a full body trait.

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Y-link Half Black (NiII), courtesy A. S. Bias

In further example of pigment repulsion, we have the new “Sunburst Swallow” phenotype of Asian Breeder Akrawat. While rather fluid in makeup and expression, it relies on co-expression of NiII + red & yellow color pigments. Though he often adds Schimmelpennig Platinum (Sc)3and Ssb traits to produce further variation. As in the Sunset Micariff, while there may be minimal blending of the two color pigments, the expected result is clear delineation; a repulsion of red color pigment and black melanophores away from yellow to the exterior edges of finnage. In Sunbrust Swallow the full body trait is NiII or Moscow Blau Additional Gene (MBAG). [Note: The presence "swallow" trait and not only it's effect on finnage, but also confinement of yellow color pigment between hemirays.]

Sunburst Swallow showing red pigment repulsion in both males and females, courtesy Akrawat Farms.

Now we are to a point where it is prudent to reveal the missing trait found in Micariff I have been alluding to. Leucophore White (Le) has been around for a number of years, though seemly overlooked by many. Yet is found in genotype of many solid colored strains and frowned upon for hindering additional color pigment coverage. In example the following photos with minimal expression similar to wild-type.

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Albino Full Red, courtesy of Paulo H. Keijock Muiz

Expanded expression of Le has been known by long-time IFGA breeders as "Parrish White" in the anterior shoulders. In his writing's Phil Shaddock refers to as leucophore body (Le) and leucophore fins (Lef). While independent regulation between body and finnage has long been accepted in general, I am no longer sure this is a valid assumption in general. For some time I have been referring to this trait as “Micariff White” (hereafter Leucophore White) in the body while in correspondence with fellow breeders in discussing the Micariff strain. Here is where I will step out on a limb and hope it does not break. To my knowledge not a single "solid" Le body and finnage strain has ever been created that does not also incorporate at least one sex-linked or autosomal "full body" trait. [Note: White guppies are normally a combination of; 1. NiII + Le. 2. American Pink White (Pw) and one or more or the following Ivory (Iv) or Asian Blau (Ab) or Sc or s]. Rather than devote extensive space to speculation and further explanation to Le qualities will simply state; Leucophore White pattern is likely the result of genetic switching in utero. Possibly it is a form of conversion between melanophores and leucophores. As can be seen in the following IFGA examples, modification of Le often appears limited to expression in the anterior body. Each of the three males lacks a visible full body trait. IFGA breeders have only been marginally successful in masking anterior Le with inclusion of iridophores by Viridis (Vir), i.e. "blue belly" trait, Full Red body traits or Mg to produce a green cast in the anterior belly / shoulder region.

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(clockwiset) IFGA Blue, Purple and Green deltas, courtesy of Bryan Chin.

In this example an IFGA Gold Delta expresses finnage comprised solely of Le, less addition of Mg and Variegation (Var). It is a true solid white fin strain encompassing at least two full body traits; b and Pb. Leucophore White is present anteriorly, incompletely masked by Mg and Pb. I like to view Mg as an opportunist; It randomly seeks out regions lacking color pigment from fixed traits. It is amplified by full body traits, to populate large areas in regions comprised of iridophores or modified leucophores.

Gold Delta, courtesy Bryan Chin

In this example a Half Black White (IFGA HB AOC) phenotype is comprised solely of Le in finnage. Or is it limited to finnage? It is a true solid white fin strain encompassing a full body trait; NiII. While white coloration of scales on the topline is often simply equated to "platina", it is the result of Le in co-expression with NiII. Platina is easily modified to red, yellow or blue.

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Half Black White, courtesy of Ramona Osche

The presence of iridophores or leucophores is required to maximize the Mg population and expression. The presence of leucophores is required for maximum yellow color pigment population and expression. The majority of "yellow" found in the bodies and finnage of guppies is the product of Mg. Sex-linked yellow pigment is found in finnage with or without Mg. Few "true" yellow color pigment body traits have been identified in guppies. The two most common in the body are Winge's Vitellinus (Vi) & Cinnamomeus (Ci). They may be found either stand alone or together in various phenotypes. To the HB White phenotype we add yellow color pigment & Mg to produce the HB Yellow. [Note: Autosomal Mg, considered a flaw in HB White, has proven hard to eliminate in many strains.]

Half Black Yellow, courtesy Bryan Chin

Probably the best known example of NiII + Le occurs in the phenotype known as the Albino Full Platinum White. It also encompasses two full body traits; NiII and a.

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Albino Full Platinum White male, courtesy Joe Putta

To this we add Mg and yellow color pigment to create the Full Platinum Gold; NiII + Le + Mg. As in the full white version the addition of a full body trait amplifies Le expression to increase Mg expression. Unlike the full white version lack of pigmentation over Le is more visible on the anterior forehead. Often as a result of no "melanophore pattern" being present near the forehead. Still much variation can be found between individuals in amount of Mg coverage near the shoulders based on traits they contain in their genotype.

Full Platinum Gold, courtesy Jarawee Guppy

The final example of interactions between melanophore based pattern and Le expression will briefly touch on Half Tuxedo (Ht), also known as Saddleback. The normal expression for Ht is black and white, as shown in the photo below. Expression of yellow is simple the inclusion of Winge's Vi &/or Ci traits, often amplified by Ht. This demonstrates that Le is not impervious to overlay by a true yellow color pigment trait.

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Vitellinus (Vi), Winge, O. (1927)

Cinnamomeus (Ci), Winge, O. (1927)

Cinnamomeus (Ci), courtesy Ronan Boutot

Cinnamomeus (Ci) + Saddleback (Ht) expressing White Leucophore (Le)

(Left photo) Vitellinus (Vi) + Saddleback (Ht),courtesy Tobias Bernsee (Right photo) Vitellinus (Vi) + Cinnamomeus (Ci) + Saddleback (Ht), courtesy Akrawat Farm

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Vitellinus (Vi) + Cinnamomeus (Ci) + Saddleback (Ht), courtesy Akrawat Farm

Leucophore White expression in females, as a distinct pattern trait is harder to gauge. The normal wild-type thoratic (chest) cavity in guppies is lined with a serous membrane with a predominant population of white leucophores. Silver iridophores, while present are less visible on the exterior lining, and more so on the interior of the lining. The translucent nature of skin structure in Guppies and reduced melanophores, in the form of reticulation, allows for visibility "beneath the skin". This is also a wild-type trait in response to predation; from below a white belly is less noticeable against the sky. From above a dark topline is less visible against the bottom. Notice the abdomens of the following females. [NOTE: A serous membrane has a more complex histology than just leucophores.]

IFGA Purple delta female, courtesy A.S. Bias

Yet we know Le does express in the finnage and body of females possessing a full body trait such as NiII, as found in the three following examples.

Albino Full Platinum White male, courtesy Joe Putta

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HB White female, courtesy Benson Liu

Half Black Blue female, courtesy Joe Putta

I first started to take interest in Le during reciprocal outcrosses involving Micariff going back nearly 15 years. In an outcross several years ago involving a Yellow Micariff male x Vienna Lowersword females the expected Ssb was produced in F1. Of particular interest note that Le remains present in pattern coexpression; Le + Ssb. Also note the extent of coverage. Not only is Le is an integral part of the Micariff phenotype. Without its presence in genotype the Micariff would not exist.

F1 Micariff x Vienna Lowersword, courtesy A.S. Bias

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In many strains a leucophore pattern is partially masked. We are only provided a glimpse of it's full expression. In Joe Putta's Albino Cherry Pink Grass strain we are afforded a view of the extended pattern coverage with melanophores eliminated, and expressing Le. [Note: Albino guppy mutants eliminate melanin rather than reduce the size or number of melanophores. Blond will reduce the size and number or melanophores.]

Albino Cherry Pink Grass, courtesy Joe Putta

Grey Cherry Pink Grass, courtesy Joe Putta

Some years back Russian breeder Vladimir Storozchev produced an intriguing phenotype known as “Marlboro Guppy” that lacks Mg. While this phenotype is comprised of numerous traits, the one that stands out in particular is Le. Here also is one of the few phenotypes to expose the extent of a Le pattern without also incorporating Mg, as found in the Micariff.

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Marlboro Guppy, bred by Vladimir Storozchev

In the KOI phenotype we see an above average expression of Le. Normally most Le is masked, except for the lower abdoman, as found in the parental Full Red strain. Full Red being comprised of a mixture of both full body and localized red traits. All of which are not found in the KOI, though males typically still express more anterior shoulder or peduncle red. To which breeders are continually infusing additional red traits from other red strains and producing variants on the initial KOI expression.

KOI females, courtesy of David Teng

Over the last several years I have been discussing and following the breeding’s of Carl Groenewegen’s X-linked Schimmelpennig Platinum’s; i.e. Royal Platinum’s. His Royal Platinum’s were started from a foundation male obtained at a local fish store. Pedigree unknown, he was a very iridescent male expressing NiII. When bred to Carl’s linebred blond red tail females he produced the F1 the male below. Hidden beneath most of his color and pattern was a trait almost overlooked, Leucophore White.

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F1 male, courtesy Carl Groenewegen

Notice the white expressing in his belly and towards the peduncle. Also white platina in topline scales and those at the peduncle base. This F1 male expresses NiII + Le + Sc. If I did not know his descendants, might think him simply a Full Platinum (NiII + Mg) in grey body. Sc expression is not limited to shoulder pattern, but has full body effect into finnage. It amplifies Mg and Le, as does NiII. Intially masked by NiII, Carl was able to isolate out Ssb, s, and Mg in later breeding’s in co-expression with Sc as X &/or Y-link. In each case Le is also visible to varying degrees. Examples of each below.

All photos courtesy Carl Groenewegen

With continued breeding’s Le was also isolated in conjunction with Sc, and improved upon to produce the Royal Platinum males below. I find the Royal Platinum phenotype to be unique not only in its combination; X &/or Y-linkSc + Le, but also the overall degree of Le expression amplified by Sc. I have not seen this combination from other breeders. Though, s + Le in co-expression are fairly common.

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Royal Platinum strain in co-expression with Leucophore White, courtesy Carl Groenewegen

Though he appears to be non-metallic and heterozgyous for Stoerzbach, after multiple generations when required traits were recombined, a Micariff Sunset was produced.

Micariff reconstructed from stand alone traits, by Carl Groenwegen

Summary: To restate my hypothesis: Mg, s, Ssb all in homozygous state, and a “missing trait” are required to produce the original metallic Micariff phenotype in full body expression. That this missing trait is required in conjunction with Steorzbach (s) a "full body" trait acting as a precursor to effect metallic Micariff phenotype in both body and finnage. As a result two actual expressions are commonly produced when Stoerzbach is lacking or in heterozygous form; metallic and non-metallic. The Micariff phenotype is very similar to the Schimmelpennig Platinum phenotype. Each has distinct zones of Le regulation, yet overall both are a silver / blue iridophore full body trait. Lacking any one of these traits (Mg, s, Ssb ) may produce a similar expression, but at best is of “Micariff” type and not truly a metallic Micariff as originally created on the island of Sri Lanka. Analysis of photos and breeder records, while not definitive, does lend credence. Le and s are both integral parts of Micariff composition. Le is not simply a trait resistant to pigmentation and a hindrance to full yellow coloration. [Note: The last observation can be made for the Full Gold Platinum phenotype.] Page 19 of 21

To prove this supposition conclusive, two approaches could be taken: 1. Reciprocal outcrosses using a pedigree Micariff strain, followed by further sib-breeding, backcrosses, and outcrosses. The resulting data could analyzed and the needed ratio’s obtained to verify composition. 2. Utilize the suggested stand-alone traits comprising pedigree Micariff as found in random stocks. Then attempt to recreate the Micariff phenotype over multiple generations, as did Carl Groenwegen, with documentation. Steorzbach (s) and Metal Gold (Mg) are autosomal and thus recombine independently from the sex-linked genes. It seems likely that they also recombine independently from each other as well. Ssb, Le and NiII are all sex linked genes. Recombination frequencies between them may allow mapping to be done, but they will each recombine independently with s and Mg. The most efficient method for separating the three sex linked genes from each other and thereby demonstrating their independent existence would be by reciprocal crossing Micaraff to a strain that lacks all three of these genes, and then test crossing the F1 back to the test strain lacking these genes. This would allow the mapping of the three genes as well as showing their presence. If the reciprocal cross uses a strain that lacks all 5 genes, that would be even better, but probably not necessary. In either case, it would require extensive use of tanks and time…

Acknowledgments Richard Squire, Ph. D. (Genetics), Retired Full Professor of Biology, University of Puerto Rico, Mayaguez Campus. For his suggestions, technical edits and proof reading of this article. Foot Notes: 1. Stoerzbach described by Kempkes (2007) as autosomal recessive Metallicus (Me). 2. Japan Blue described by Kempkes (2007) as Y-link Caeruleus (Ca). 3. Schimmelpennig Platinum described by Kempkes (2006) described as Y-link Buxeus ( ). No putative notation suggested by author resulting from incomplete breeding results.

Genetic Notations: Albino (a) American Pink White (Pw) Asian Blau (Ab) Blond (b) Cinnamomeus (Ci) Full Gold (NiII + Mg) Half Black (NiII) Japan Blue / Aquamarine (A) / Caeruleaus (Ca) Lazuli (Lz) Leucophore White (Le) Metal Gold (Mg) Moscow Blau Additional Gene (MBAG) American Pink White (Pw) Ivory (Iv) Purple Body Mutation (Pb) Schimmelpennig Platinum (Sc) / Buxeus ( ) Snakeskin Body (Ssb) Snakeskin Tail (Sst) Stoerzbach (s) / Metallicus (Me) Variegation (Var) Viridis (Vir) Vitellinus (Vi)

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References: Arnheim, R. (1969), Visual thinking. Berkeley; Univ of CA Press. BAGNARA JT (1958), Hypophyseal control of guanophores in anuran larvae. J Exp Zool. 1958 Mar;137(2):265–283. Bagnara, JT (1968), The Dermal Chromatophore Unit, J Cell Biol., v. 38(1); 1968 Jul 1, PMC2107474. Grasseni, C (2004), Skilled vision. An apprenticeship in breeding Aesthetics, Social Anthropology (2004), 12, 1, pgs. 41-55. Grasseni, C (2005), Designer Cow: The Practice of Cattle Breeding Between Skill and Standardization, Society & Animals, 13:1. Grether, G (2004), Individual colour patches as multi component signals, Biol. Rev. (2004), 79, pp. 583-610. Kempkes, M (2007), New colour genes in the guppy, Poecilia reticulate (Peters, 1859), Bulletin of Fish Biology, Vol. 9, Nos. 1 / 2, 15.12.2007, pgs. 93-97. Kempkes, M (2006), Buxeus, a new colour in the guppy, Poecilia reticulate Peters, 1859 (Teleostei, Cyprinodontiformes), Zeitschrift fur Fischkunde, Vol. 8, Nos. 1 / 2, 15.10.2006, pgs. 99-100. Shaddock, P (2012), Guppy Mutations Under the Microscope, August, 2012, ISBN: 978-0-986570-2-5, Published by Pocket Cine, Inc. Yellow Guppy, The Online Home of American Guppy Breeder Mike Khalid, http://yellowguppy.com/. 3.24.15. Note: Some photo links broken.

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