this was subsequently made a separate genus. Diplacus is clearly derived from within Mimulus, broadly defined, and was not usually considered to be a separate genus. The French botanist Édouard Spach established Erythranthe as a separate genus with just the type species Erythranthe cardinalis. In 1885, American botanist Edward Lee Greene classified Erythranthe as a section of Mimulus while adding E. lewisii and E. parishii. In the 2012 restructuring of Mimulus by Barker et al., based largely upon DNA evidence, seven species were left in Mimulus as strictly defined; Erythranthe was greatly enlarged to include 111 species, based upon axile placentation and long pedicels, 46 placed into Diplacus (species with parietal placentation and sessile flowers), two placed in Uvedalia, and one each placed in Elacholoma, Mimetanthe, and Thyridia. All of the American genera are still referred to as "monkey-flowers". Views on the evolutionary position of the monkey-flower species have changed. It was long considered to be in the family Scrophulariaceae, but is now placed in Phyrmaceae, primarily on the basis of DNA evidence. The genus Phryma (comprising only a single species), for which the family is named, is considerably different in morphology from all of the monkey-flowers. Attempts at crossing species, whether from different sections or within the same section, of Erythranthe are not always successful. E. peregrina is an example of a successful naturally occurring hybrid that not only arose independently in two different locations, but is also a rare example of evolutionary recent allopolyploidization, complete chromosomal inheritance. Charles Darwin's 1876 study of inbreeding depression and self-fertility in South American species was a progenitor for the study of Erythranthe biology. The genus has become a model system "for studies of evolutionary and ecological functional genomics ... [as it] ... contains a wide array of phenotypic, ecological and genomic diversity." Species under intense genomic study are mostly among the section Simiolus (E. guttata and relatives) and the section Erythranthe (including E. lewisii, E. cardinalis, E. parishii, and others). The genome sequence of E. guttata was released in late spring, 2007. Many issues remain in Erythranthe taxonomy. E. guttata is highly complex, with many variations apparently reflecting differences in geographic environment and elevation. Molecular geneticists regard the species broadly as including both perennial and annual populations, but there is rationale for treating this complex as several distinct species (perennials are E. guttata, E. grandis, and E. corallina; annuals are E. microphylla