Nucleomorphs are small, vestigial eukaryotic nuclei found between the inner and outer pairs of membranes in certain plastids. They are thought to be vestiges of primitive red and green algal nuclei that were engulfed by a larger eukaryote. Because the nucleomorph lies between two sets of membranes, nucleomorphs support the endosymbiotic theory and are evidence that the plastids containing them are complex plastids. Having two sets of membranes indicate that the plastid, a prokaryote, was engulfed by a eukaryote, an alga, which was then engulfed by another eukaryote, the host cell, making the plastid an example of secondary endosymbiosis. Organisms with known nucleomorphs So far, only two monophyletic groups of organisms are known to contain plastids with a vestigial nucleus or nucleomorph: the cryptomonads of the supergroup Chromista and the chlorarachniophytes of the supergroup Rhizaria, both of which have examples of sequenced nucleomorph genomes. Studies of the genomic organization and of the molecular phylogeny have shown that the nucleomorph of the cryptomonads used to be the nucleus of a red alga, whereas the nucleomorph of the chlorarchniophytes was the nucleus of a green alga. In both groups of organisms the plastids originate from engulfed photoautotrophic eukaryotes. Of the two known plastids that contain nucleomorphs, both have four membranes, the nucleomorph residing in the periplastidial compartment, evidence of being engulfed by a eukaryote through phagocytosis. In addition, some species within the dinoflagellates that have gone through tertiary endosymbiosis also have endosymbionts with both a nucleus and mitochondria present. Nucleomorph genome Nucleomorphs represent some of the smallest genomes ever sequenced. After the red or green alga was engulfed by a cryptomonad or chlorarachniophyte, respectively, its genome was reduced. The nucleomorph genomes of both cryptomonads and chlorarachniophytes converged upon a similar size from larger genomes. They retained only three chromosomes and many genes were transferred to the nucleus of the host cell, while others were lost entirely. Chlorarachniophytes contain a nucleomorph genome that is diploid and cryptomonads contain a nucleomorph genome that is tetraploid. The unique combination of host cell and complex plastid results in cells with four genomes: two prokaryotic genomes (mitochondrion and plastid of the red or green algae) and two eukaryotic genomes (nucleus of host cell and nucleomorph). The model cryptomonad Guillardia theta became an important focus for scientists studying nucleomorphs. Its complete nucleomorph sequence was published in 2001, coming in at 551 Kbp. The G. theta sequence gave insight