Fumariaceae DC.

Including Chylaceae Dulac

Excluding Hypecoaceae, Pteridophyllaceae

Habit and leaf form. Herbs. Perennial; with a basal aggregation of leaves; rhizomatous, or tuberous. Climbing (sometimes scandent), or self supporting; when scandent, climbing via modified petiolules. Mesophytic. Leaves alternate (to sub-opposite); spiral; petiolate; non-sheathing; simple, or compound; when compound, ternate, or pinnate, or bipinnate, or multiply compound. Lamina when simple, usually dissected; when simple, pinnatifid, or palmately lobed, or much-divided; pinnately veined. Leaves exstipulate; without a persistent basal meristem.

Leaf anatomy. Minor leaf veins without phloem transfer cells (Corydalis, Dicentra, Fumaria).

Stem anatomy. Nodes unilacunar (usually), or tri-lacunar. Secondary thickening absent (?). Xylem with vessels. Vessel end-walls simple. Sieve-tube plastids S-type.

Reproductive type, pollination. Hermaphrodite.

Inflorescence, floral, fruit and seed morphology. Flowers solitary (rarely), or aggregated in `inflorescences' (usually); when aggregated, in racemes (i.e. usually). The terminal inflorescence unit cymose. Inflorescences `usually more or less racemose'. Flowers very irregular; zygomorphic; 2 merous; cyclic. Free hypanthium absent.

Perianth with distinct calyx and corolla; 6; 3 whorled; isomerous. Calyx 2; 1 whorled; polysepalous; not persistent (caducous, scalelike); open in bud. Corolla 4; 2 whorled; polypetalous to gamopetalous (more or less connivent, the two inner members more or less coherent over the stigmas apically); imbricate; spurred (or at least saccate, basally, in terms of one or both members of the outer whorl).

Androecium 6. Androecial members free of the perianth; coherent; 2 - adelphous (i.e. in two bundles of three, the bundles opposite the outer corolla members). Androecium exclusively of fertile stamens. Stamens 6. Filaments appendiculate (with basal nectaries), or not appendiculate. Anthers extrorse; unilocular (the lateral members of each triplet), or bilocular (the central member - i.e. the stamens dimorphic within each triplet); bisporangiate and tetrasporangiate. Endothecium developing fibrous thickenings. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral, or decussate. Anther wall initially with one middle layer; of the `monocot' type. Pollen grains aperturate; 3 - aperturate, or 6-12 - aperturate; colpate, or rugate (tricolpate or 6-12 rugate); 2-celled.

Gynoecium 2. Carpels isomerous with the perianth. Gynoecium syncarpous; eu-syncarpous; superior. Ovary 1 locular. Gynoecium transverse; stylate. Styles 1; apical. Stigmas dorsal to the carpels, or dorsal to the carpels and commissural; capitate. Placentation parietal. Ovules in the single cavity 2-100 (to `many'); arillate; anatropous to campylotropous; bitegmic; crassinucellate. Outer integument contributing to the micropyle. Embryo-sac development Polygonum-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; not proliferating; very large. Synergids at least sometimes with filiform apparatus. Endosperm formation nuclear. Embryogeny caryophyllad.

Fruit non-fleshy; dehiscent (usually), or indehiscent (rarely); a capsule, or a silicule, or a siliqua, or a nut (rarely). Capsules loculicidal, or valvular (or breaking transversely into 1-seeded segments). Fruit 1-100 seeded. Seeds endospermic. Endosperm oily. Embryo well differentiated (small). Cotyledons 1 (e.g. some Corydalis spp.), or 2. Embryo achlorophyllous (2/2); straight to curved.

Physiology, biochemistry. Not cyanogenic. Alkaloids present. Iridoids absent. Proanthocyanidins absent. Flavonols present; kaempferol, or kaempferol and quercetin. Ellagic acid absent (2 genera, 2 species).

Geography, cytology. Temperate to sub-tropical. Widespread North temperate, a few in montane Southern and Eastern Africa. X = (6-)8.

Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgren's Superorder Ranunculiflorae; Ranunculales. Cronquist's Subclass Magnoliidae; Papaverales. Takhtajan's Subclass Ranunculidae; Ranunculanae; Papaverales. Species 450. Genera 15; Adlumia, Capnoides, Ceratocapnos, Corydalis, Cryptocapnos, Cysticapnos, Dactylicapnos, Dicentra, Discocapnos, Fumaria, Platycapnos, Pseudofumaria, Rupicapnos, Sarcocapnos, Trigonocapnos.

Illustrations. fumar219.gif fumar220.gif

Additional, to be intercalated. Flowers when solitary, axillary.