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 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.

Before 2020, only two monophyletic groups of organisms were known to contain plastids with a vestigial nucleus or nucleomorph: the cryptomonads of the supergroup Cryptista 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. Both have four membranes, the nucleomorph residing in the periplastidial compartment, evidence of being engulfed by a eukaryote through phagocytosis.

In 2020, genetic work identified the plastid in Lepidodinium and two previously undescribed dinoflagellates ("MGD" and "TGD") as being most closely related to the green alga Pedinomonas. The observation of a nucleomorph in Lepidodinium is controversial, but MGD and TGD are proven to have DNA-containing nucleomorphs. The transcriptomes of the nucleomorphs have been sequenced. One slight issue in understanding the sequence of evolution is that although the phylogenetic tree built from Lepidodinium-MGD-TGD's plastid is monophyletic, the tree built from their host-nucleus DNA is not, implying that they might have acquired very similar algae independently.

A cryptomonad nucleomorph is typically much smaller than the host nucleus. A relatively large portion of its size is devoted to the nucleolus, which contains its own ribosomes and rRNA. There seems to be nuclear pores observable by imaging, but genetic work has failed to find any protein appropriate for forming the nuclear pore complex.

There is one nucleomorph per plastid. The nucleomorph divides before the accompanying plastid. The dividing nucleomorph lacks a mitotic spindle, and the nucleomorph envelope persists throughout division.

Between the plastid and the cytoplasm of the host there are four membranes: the inner and outer membranes of the chloroplast, the periplastid membrane, and the epiplastid membrane. The epiplastid membrane is encrusted with ribosomes (in cryptomonads) and is in many ways similar to a endoplasmic reticulum, hence the name "chloroplast endoplasmic reticulum" (cER). Plastid-targeted proteins encoded in the host genome must cross all four membranes to reach the plastid. First they use classic secretory signal peptides to cross the epiplastid membrane. Then the symbiont-specific ERAD-like machinery (SELMA) – encoded in the nucleomorph as a repurposed ERAD – pulls the protein from the epiplastid space (or the lumen of the cER) into the periplastid space (the cytoplasm of the symbiote). The standard chloroplast transit peptide then acts to cross the remaining two layers via TIC/TOC complex.

The chlorarachniophytes, on the other hand, has no such thing as a cER, hence the initial import into the epiplastid space must occur by some other mechanism. It's only known that their plastid-targeted proteins are prefixed by both a signal peptide and a chloroplast-targeting peptide much like cryptomonads. Based on research done on apicomplexa, which also has 4 membranes but no cER, it's possible that the protein is first sent into the ER, then sent to the epiplastid space by the endomembrane sorting system. Some sort of a pore may then move the peptide into the periplastid space, but there seems to be no SELMA-like pore in this group. It's only known that the TIC/TOC complex exists for crossing the last two layers.

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).