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POECILIID RESEARCH International Journal of the Bioflux Society- Research Article POECILIID RESEARCH International Journal of the Bioflux Society- Research Article A search for origins of iridescence in modern domesticA guppies; if and when did introgression occur? Alan S. Bias Swordtail Guppy Breeder & Designer, P.O. Box 1508, Lewisburg, West Virginia 24901, USA. Corresponding author: A. S. Bias, [email protected] Abstract. The source of genetic inputs in modern Domestic2 Guppy strains has long been a controversial topic in breeder circles. Assumed by many to consist solely of Poecilia reticulata, and considered by others to have multiple inputs from closely related variant populationsB. The attempt of this paper is to demonstrate that sum genotype, in particular for at least one identifiable trait known as Iridescens (Ir), stems from an amalgamation through interbreeding of native GuppyC populations in divergent stages of incipient speciation6; that are capable of interbreeding and surviving in either habitat, and are therefore non-cryptic1. Offspring should not be viewed as interspecific hybrids5 (between different species in the same genus), as they are capable of viable reproduction, lacking any high degree of decrease in reproductive viability. Domestic strains, like wild counterparts in reproductive isolation11 are not speciated. Results are based on 45 years breeding experience of which nearly thirty (30) years have been devoted to strains collectively known as “Swordtail Guppies”, efforts of other professional breeders, and that of the scientific community. Key Words: swordtail guppy, domestic strains, Vienna emerald, wild-type, guppy populations, reflective qualities, smaragd-iridescens, iridescence, Poecilia reticulata, Poecilia obscura, Poecilia wingei, Cumana’ guppy, Campoma guppy, Endler’s livebearer. Hypothesis. Based on what we know today, it is likely early 1900’s importations of the Guppy (Poecilia reticulata) made available to breeders contained “iridescent-reflective” genotype (Photo 1), as a result of admix of variant populations. Therefore, future introductions and usage of variants, by breeders, in domestic stocks should be viewed simply as “genetic tools” for improvement. Does phenotypical evidence exist to support admix? Photo 1. Iridescent Asian Blau Schimmelpennig Platinum (Domestic Strain) (photo by A. S. Bias). Poeciliid Research, 2013, Volume 3, Issue 1. http://www.pr.bioflux.com.ro/ 22 Introduction. What defines a species12 from a related variety or population? Species; Quote: “Species are groups of actually or potentially interbreeding natural populations, which are reproductively isolated from other such groups” (Mayr 1942). Mayr formulated this statement as part of his belief in the concept of Biological Species (Dobzhansky 1935, 1950) which in brief states members of species can only breed among themselves. This means since crosses between members of different populations produce fertile F1 and F2 offsprings, they cannot be considered interspecific crosses as they would fail to produce an F2. This concept is based on the assumption that while geographic isolation plays a large part in speciation, so does genetics in the form of an Isolating Mechanism7. The existence of the Guppy has been known to science since 1859 when German Zoologist Wilhelm C. H. Peters first published results based on preserved specimens collected in 1856 from the Guayre River outside of Caracas, Venezuela. This moment of stability would be lost for the next 50 plus years as P. reticulata would be re-classified and confused not only with other populations, but its own. DNA results based on a single specimen taken from the 1856 collection suggest: “This sequence clusters with all present-day P. reticulata (Photo 2) including a fish taken in 2008 from a tributary to the Guayre…” (Schories et al 2009). Photo 2. Wild-type P. reticulata, Brazil collection (photo courtesy of Shimpei Taniguchi). Prior to 1910 there is little or no reference to the existence of Guppies in the realm of emerging stock breeders. During the years 1905-14 German breeders were just starting to develop their interest and means to collect and/or import wild guppies. Whether these live specimens came from Trinidad or mainland South America cannot be ascertained with any degree of certainty. This is an important issue, as later it will be demonstrated that one location plays a larger role in potential source of iridescent qualities than the other. It is believed that all North American stocks were obtained from German breeders until the time of World War I. During the period of 1918-1927 in a series of studies and resulting research papers, Prof. Ö. J. Winge of Copenhagen Denmark, published his findings involving the genetics of numerous Guppy phenotypes. While color photography was in its infancy and apparently not used to document results, we are fortunate to have accurate color representations of his findings. One trait in particular (Photo 3), Iridescens (Ir), is of particular interest for further discussion at a later point in this writing. Poeciliid Research, 2013, Volume 3, Issue 1. http://www.pr.bioflux.com.ro/ 23 Photo 3. Winge (1927), Illustrations of Y-link gene for Iridescens (Ir). Definitions: 1. Cryptic species complex: similar on a phenotypic level, but DNA testing reveals genetically distinct. Interbreeding does not occur, otherwise there would be loss of distinction. 2. Domestication: to bring of keep (wild animals or plants) under control or cultivation (http://dictionary.reference.com). 3. Founders: individuals breeding members, initially responsible for founding a breeding population. 4. Genetic drift: gene frequencies within strains change over time due to random events by chance alone; resulting in a possible loss of certain traits. In small populations selection is often skewed and beneficial alleles may be lost. 5. Hybrid: (Life Sciences & Allied Applications/Biology) an animal or plant resulting from a cross between genetically unlike individuals. Hybrids between different species are usually sterile (http://www.thefreedictionary.com). 6. Incipient speciation: populations that are in the process of diverging to the point of speciation but still have the potential to interbreed (http://www.answers.com). 7. Isolating mechanism: something (as a geographical, ecological, physiological, anatomical, or psychological barrier) that limits interbreeding gene flow between groups and is thereby a major factor in the differentiation of biologic units (as races or species) (http://www.merriam-webster.com). 8. Mutation: a mutation is a permanent change in the DNA sequence of a gene. Mutations in a gene's DNA sequence can alter the amino acid sequence of the protein encoded by the gene (http://learn.genetics.utah.edu/archive/mutations). 9. Natural selection: the process by which plants and animals that can adapt to changes in their environment are able to survive and reproduce while those that cannot adapt do not survive (http://www.merriam-webster.com) 10. Phylogeny: the evolutionary development and history of a species or higher taxonomic grouping of organisms (http://www.thefreedictionary.com). 11. Reproductive isolation: the inability of a species to breed successfully with related species due to geographic, behavioral, physiological, or genetic barriers or differences (http://www.merriam-webster.com). 12. Species: a fundamental category of taxonomic classification, ranking below a genus or subgenus and consisting of related organisms capable of interbreeding (http://www.thefreedictionary.com). Discussion Identification of wild-type Guppy populations. Part of the problem in classification of variants comprising Acanthophacelus (Eigenmann 1907) has been compounded by the inability of science to determine just what composes a common Guppy for lack of an accurate phylogeny10. Science has not provided a singular description entailing natural ranges, variation in color patterns, anatomical, behavioral variations, and historical processes. Are variant populations truly incipient or cryptic? They freely interbreed with little or no evidence of lethal heterozygous alleles for environmental or hereditary conditions in Poeciliid Research, 2013, Volume 3, Issue 1. http://www.pr.bioflux.com.ro/ 24 offspring. Little in the way of MtDNA differentiation (Breden Labs) has been noted. Together this suggests incipiency. If geographic barriers on mainland South America once existed, lack of genetic distance in variants (P. reticulata & Poecilia wingei) indicates any prior genetic differentiation towards speciation may be in the process of recombination (Webb et al 2011). Support can be found in Schories et al (2009), quote: We hypothesize that P. wingei was the most western species but recently was circled by northwest colonizing populations of P. reticulata. We are left with sexual selection as the primary driving force behind variation in populations. While the debate is ongoing and will likely be so for some years to come, in the last decade some clarification has been provided by science for the benefit of breeders. Findings suggest that the sub-genus Acanthophacelus (Eigenmann 1907) currently has at minimum three or more known variant populations: 1. The Orinoco (Common) Guppy; P. reticulata (Peters 1859)E, 2. The Endlers/Cumana’/Campoma Guppy; P. wingei n. sp. (Poeser et al 2005), and 3. The Oropuche Guppy; Poecilia obscura n. sp. (Schories et al 2009). In his description of P. wingeiG as a distinct species Poeser et al (2005) states, quote: “Its closest relative is the common guppy, P. reticulata, sharing identical meristic
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