The families of flowering plants

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L. Watson and M. J. Dallwitz

Fumariaceae DC.

~ Papaveraceae Juss.

Including Chylaceae Dulac; excluding Hypecoaceae, Pteridophyllaceae.

Habit and leaf form. Herbs; laticiferous (the latex watery). Annual, or perennial; with a basal aggregation of leaves, or without conspicuous aggregations 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 palmatifid, or much-divided; pinnately veined. Leaves exstipulate; leaf development not ‘graminaceous’.

General anatomy. Plants with laticifers.

Leaf anatomy. Hairs present, or absent; unicellular. Minor leaf veins without phloem transfer cells (Corydalis, Dicentra, Fumaria).

Axial (stem, wood) anatomy. Young stems commonly with hollow internodes. Nodes unilacunar (usually), or tri-lacunar. Primary vascular tissues comprising a ring of bundles; collateral. Cortical bundles absent. Medullary bundles absent. Secondary thickening absent, or absent to developing from a conventional cambial ring (?). Primary medullary rays wide.

The vessel end-walls simple.

Reproductive type, pollination. Fertile flowers hermaphrodite. Plants hermaphrodite.

Inflorescence, floral, fruit and seed morphology. Flowers solitary (rarely), or aggregated in ‘inflorescences’ (usually); when solitary, axillary; when aggregated, in racemes (i.e. usually). The ultimate inflorescence units racemose. Inflorescences terminal (?), or axillary, or leaf-opposed (commonly); ‘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; neither appendaged nor spurred (the sepals not lobed); not persistent (caducous, scalelike); open in bud (not enclosing the flower bud). Corolla 4; 2 whorled (2+2); 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 ostensibly 6. Androecial members branched (in that the two lateral units mostly consist each of one dithecal and two monothecal units); 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 carpelled. Carpels isomerous with the perianth. The pistil 1 celled. 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 (usually with two placentas). Ovules in the single cavity 1 (sometimes, in Fumaria), or 2–100 (each placenta with ‘one 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), or lomentaceous; a capsule, or a silicula, or a siliqua, or a nut (rarely). Capsules loculicidal, or valvular (or breaking transversely into 1-seeded segments). Fruit 1–100 seeded (i.e. to ‘many’). 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, phytochemistry. Not cyanogenic. Alkaloids present. Iridoids not detected. Proanthocyanidins absent. Flavonols present; kaempferol, or kaempferol and quercetin. Ellagic acid absent (2 genera, 2 species). Sieve-tube plastids S-type.

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. APG 3 core angiosperms; peripheral eudicot; Superorder Ranunculanae; Order Ranunculales (as a synonym of Papaveraceae).

Species 450. Genera 15; Adlumia, Capnoides, Ceratocapnos, Corydalis, Cryptocapnos, Cysticapnos, Dactylicapnos, Dicentra, Discocapnos, Fumaria, Platycapnos, Pseudofumaria, Rupicapnos, Sarcocapnos, Trigonocapnos.

General remarks. The data compiled for this package exhibit conspicuous differences between Fumariaceae and Papaveraceae (q.v.) involving seven morphological characters; however, the taxonomic justification for retaining the separate families is reduced if the current descriptions significantly under-estimate intra-familial variation. The anatomical compilation of Metcalfe and Chalk for Papaveraceae sensu lato and Fumariaceae is not reliaby interpretable in the present context.

Illustrations. • Technical details: Corydalis, Fumaria. • Technical details: Ceratocapnos, Cysticapnos, Dicentra. • Corydalis solida and Corydalis (= Pseudofumaria) lutea: Eng. Bot. 68 and 69, 1863. • Fumaria bastardii (as F. confusa): Eng. Bot. 73, 1863. • Fumaria densiflora (as F. micrantha), F. officinalis, F. parviflora: Eng. Bot. 75, 76 and 78, 1863. • Fumaria muralis ssp. boraei: Eng. Bot. 72, 1863. • Corydalis, Fumaria (B. Ent. compilation.


And Fumitory too, a name
Which superstition holds to fame,
Whose red and purple mottled flowers
Are dropped by maids in weeding hours,
To boil in water, milk, and whey,
For washes on a holiday,
To make their beauty fair and sleek,
And scare the tan from summer’s cheek

