Seed-coat anatomy in Fumariaceae-Fumarioideae

Seed-coat anatomy is described in 122 species of Fumarioideae, which represent all the genera, subgenera, and most sections. Nine seed-coat types were recognized: (I) Dicentra subg. Hedycapnos , (II) Dicentra subg. Chrysocapnos , (III) Dicentra subg. Macranthos , (IV) Dicentra subg. Dicentra , (V) Corydalis p.p. , (VI) rest of Corydaleae, Cysticapnos, Pseudofumaria and Ceratocapnos , (VII) Discocapnos and Sarcocapnos p. p. , (VIII) Sarcocapnos p. p., Platycapnos and Trigonocapnos , and (IX) Fumariinae. Variable characters are polarized based on the comparison with Fumariaceae Hypecooideae, Papaveraceae and Pteridophyllaceae. It is assumed that endotestal seed-coat type (I) is most primitive, that exotestal seed coat types (II—VIII) are derived therefrom, and that the reduction of the mechanical layer, including the origin of thin seed-coat type (IX) occurred repeatedly in combination with indehiscent, hard-walled fruits. Dicentra subg. Hedycapnos (with type I) possesses a combination of primitive testal characters unique within the subfamily, suggesting it represents a sister group to the rest of the subfamily. Each of the other subgenera (with II, III or IV) is also characterized by a unique combination of seed-coat characters, which are found in the outgroups but not in the other genera. Seed-coat characters support the monophyly of Fumarioideae exclusive of Dicentra subg. Hedycapnos , Fumarioideae exclusive of Dicentra , of Dicentra subg. Dicentra , of Corydalis p.p. , and of Fumariinae.