FUMARIACEAE (fumitory family)

Plants annual, biennial, or perennial herbs, glabrous (except for the fruits in Corydalis crystallina). Leaves alternate or basal, lacking stipules, pinnately or pinnately then ternately 2–4 (or more) times compound (often appearing highly dissected with relatively narrow ultimate segments). Inflorescences terminal (sometimes also appearing axillary in Fumaria), short to elongate racemes, sometimes reduced to clusters, the flowers subtended by small bracts. Flowers irregularly zygomorphic (bilaterally symmetrical in only 1 longitudinal plane or in 2 perpendicular planes), hypogynous, perfect. Calyces of 2 free sepals, these similar in size and shape, inconspicuous, often shed as the flower opens. Corollas of 4 petals, these free or fused at the base, dissimilar in size and shape, positioned in 2 whorls, the inner whorl of 2 shorter petals more or less apically hooded or fused (enclosing the anthers and stigma), the outer whorl of longer petals with 1 or both having a basal pouch or spur (except in cleistogamous flowers). Stamens 6, positioned in 2 clusters of 3, each cluster opposite an outer petal, the filaments within a cluster fused, at least toward the base (and sometimes also to the petals), often with a basal, spurlike, nectar-producing outgrowth, the anthers small, attached at or near the base, often more or less fused around the stigma, only the middle anther of each trio with 2 pollen sacs, the flanking pair with 1 pollen sac. pistil 1 per flower, of 2 fused carpels. Ovary superior, 1-locular, the placentation parietal. Style 1 per flower, unbranched, persistent at fruiting (except in Fumaria), the stigma usually flattened, 2–8-lobed, the lobes sometimes appearing as horns or wings. Ovules 2 to numerous. Fruits indehiscent and nutlike or capsules that are longitudinally dehiscent by 2 valves (sometimes tardily so), 1- to numerous-seeded. Seeds (except in Fumaria) usually somewhat flattened, nearly circular to more commonly finely notched or somewhat kidney-shaped, the embryo curved, the surface dark brown to black, usually with a pale elaiosome (oil-bearing aril-like appendage attractive to ants). About 19 genera, about 450 species, North America, Europe, Asia, Africa.

The Fumariaceae are closely related to the Papaveraceae and have been included within that family (usually as a subfamily) by a number of authors (Lidén, 1986; Judd et al., 2002). Relationships within the lineage that includes both of these groups is still somewhat controversial and it is unclear whether the best approach is to recognize one morphologically diverse family or split the lineage into two or more families. Within the group, Fumariaceae are distinguished morphologically by their asymmetrical or zygomorphic corollas, clear sap, and unusual stamen and stigma morphology. Like the Papaveraceae, members of the Fumariaceae produce a large variety of isoquinoline alkaloids. This accounts for their various medicinal uses (Moerman, 1998) as well as their toxicity to livestock and humans (Burrows and Tyrl, 2001). A number of species are cultivated as ornamentals.