Seed and Funicle Morphology of Fumariaceae‐Fumarioideae: Systematic Implications and Evolutionary Patterns

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Abstract
This article reports the seed and funicle morphology of Fumarioideae with dehiscent fruits, covering 11 genera (106 species) for seeds and 11 genera (48 species) for funicles. The results are discussed referring to a hypothetical phylogeny based on chloroplast DNA sequences and morphology. Weak seed curvature is a synapomorphy of the Ichtyoselmis‐Dicentra clade, and the truncate hilar region and laterally elongated arilbase ambiguously support the Dactylicapnos‐Corydalis clade, but in both cases, it is necessary to assume reversals or parallel changes elsewhere in the subfamily. Otherwise, seed morphological characters are poor phylogenetic markers at the genus level as a result of high homoplasy and polymorphism within the operational taxonomic units. Although seed morphology is not very informative for the phylogeny among the terminal taxa, most of them can be identified by a combination of characters. Ehrendorferia, Dicentra, Dactylicapnos sect. Dactylicapnos (except for Dactylicapnos paucinervia), Cysticapnos s. str., and Pseudofumaria each possesses a unique hilar region topography and is supported as monophyletic. Three or five independent origins of arils are assumed in Fumarioideae. This is compatible with the diversity in the location of the arils (raphe vs. raphe‐hilar region) and with the morphology of the primordia. Each taxon possesses either hilar concavity or funicle receptacles, both of which may contribute to stabilization of seeds in fruits. The former state is assumed to have originated independently in four clades. A functional association is indicated among distinctly truncate hilar regions, platelike funicle apices, and explosive fruit dehiscence in Corydalis subg. Corydalis. These results emphasize the importance of funicle morphology in the functional interpretation of seed morphology.

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