Cardamine flexuosa |
Cardamine californica |
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wavy bitter-cress, wood bitter-cress, woodland bitter-cress |
California toothwort, coast toothwort, milk maids, tooth wort |
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Habit | Annuals or biennials; sparsely to densely hirsute basally or throughout, or glabrous. | Perennials; usually glabrous, rarely minutely pubescent. |
Rhizomes | absent. |
(tuberiform, fragile), globose to ovoid or suboblong, (3–)4–10 mm diam., (fleshy, deeply underground). |
Stems | erect, ascending, or decumbent, branched or unbranched, (0.6–)1–5 dm, (slightly flexuous). |
erect, unbranched, (2–)2.7–6(–7)dm, usually glabrous, rarely pubescent. |
Basal leaves | (often withered by anthesis), not rosulate, 5–15-foliolate, (2.7–)4–14(–19) cm, leaflets petiolulate; petiole 0.7–5 cm, (ciliate or not); lateral leaflet blade oblong, ovate, or elliptic, smaller than terminal, margins entire, repand, crenate, or 3 (or 5)-lobed; terminal leaflet (petiolule 0.3–1.7 cm), blade reniform, broadly ovate, or suborbicular, 0.5–2.5 cm × 4–30 mm, margins repand, crenate, or 3 or 5-lobed. |
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Rhizomal leaves | 3 (or 5–7)-foliolate, sometimes simple, 8–25(–38) cm, leaflets petiolulate or subsessile; petiole (5–)8–25(–32) cm; lateral leaflets (when present) petiolulate to subsessile, blade similar to terminal, sometimes smaller; terminal leaflet (petiolule (0.7–)2–5(–11) cm), blade (of simple leaf) ovate to orbicular to broadly cordate or reniform, (1.5–)2.5–7.5(–10) cm × (12–)20–90(–130) mm, base obtuse to cordate, margins entire or dentate to shallowly sinuate, (often with apiculae at veins ending at margin, sometimes minutely pubescent on veins). |
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Cauline leaves | 3–15, 5–15-foliolate [leaves (2–)3.5–5.5(–7) cm, including petiole], petiolate, leaflets petiolulate; petiole base not auriculate; lateral leaflets similar to basal, (0.4–2.5 mm wide). |
2–5, usually 3 (or 5)-foliolate, rarely simple, petiolate, leaflets petiolulate or sessile; petiole 1–5(–9) cm, base not auriculate; lateral leaflets sessile, blade similar to terminal, smaller, margins usually dentate, rarely entire; terminal leaflet sessile or petiolulate, blade usually broadly ovate to suborbicular or lanceolate, rarely narrowly oblong, 1–7 cm × (5–)10–47(–65) mm. |
Racemes | ebracteate. |
ebracteate. |
Flowers | sepals oblong, 1.5–2.5 × 0.7–1 mm, lateral pair not saccate basally; petals white, spatulate, 2.5–4(–5) × 1–1.7 mm; (stamens rarely 4, lateral pair absent); filaments 2–3 mm; anthers ovate, 0.3–0.5 mm. |
sepals (erect to ascending), oblong, 3.5–4.5(–5.5) × 1.5–2(–2.5) mm, lateral pair saccate basally; petals white to pale rose, often broadly obovate, 8–13(–15) × 4–8 mm (clawed, apex rounded); filaments: median pairs 4–6.5 mm, lateral pair 3–4 mm; anthers oblong, 1.2–1.7 mm. |
Fruiting pedicels | divaricate or ascending, (5–)6–14(–17) mm. |
ascending to divaricate, 10–33(–41) mm. |
Fruits | linear, (torulose), (0.8–)1.2–2.8 cm × 1–1.5 mm; ovules 18–40 per ovary; style 0.3–1(–1.5) mm. |
linear, 2.2–5.4(–6) cm × 2–3 mm; ovules 12–22 per ovary; style 2–5(–6) mm. |
Seeds | brown, oblong or subquadrate, 0.9–1.5 × 0.6–1 mm, (narrowly margined or not). |
dark brown, oblong to broadly ovoid, 1.7–2.8 × 1.2–1.8 mm. |
2n | = 32. |
= 32. |
Cardamine flexuosa |
Cardamine californica |
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Phenology | Flowering Apr–Jul. | Flowering Jan–May. |
Habitat | Disturbed areas, fields, nurseries, plantations, gardens, flower beds, lawns, roadsides | Wooded ravines, forest floors, shady slopes, open woods, shady rock crevices, stream banks and bottoms, canyons, moist hillsides, cliffs |
Elevation | 0-1100 m (0-3600 ft) | 0-1400 m (0-4600 ft) |
Distribution |
AL; CA; FL; GA; IL; IN; LA; MD; MI; NC; NY; OH; OR; RI; TX; VA; WA; BC; NF; ON; Europe; e Asia [Introduced in North America; introduced also in Mexico, Central America, South America, Australia]
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CA; Mexico (Baja California)
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Discussion | According to J. Lihová et al. (2006), the populations referred to Cardamine flexuosa in North America comprise two taxa of different polyploid origins and evolutionary histories: tetraploid C. flexuosa (2n = 32), native to Europe, and the octoploid taxon informally called “Asian C. flexuosa” (2n = 64), native to eastern Asia. For the latter, the name C. flexuosa subsp. debilis can be used. Nevertheless, these two taxa should be recognized at species level and the correct name for the Asian species should be sought. Based on available data, both taxa occupy the same habitats in North America, but the Asian taxon is much more widespread. The occurrence of European C. flexuosa was, until now, confirmed only for Washington, where both taxa have been recorded. More detailed studies of the North American distributions of both these weeds are needed. (Discussion copyrighted by Flora of North America; reprinted with permission.) |
The synonymy above suggests that Schulz, Detling, and Rollins did not agree on the characters used or the number and rank of taxa recognized. The taxonomy of this complex is based solely on differences in the number of leaflets, their division, size, and shape, all other aspects of these plants (e.g., rhizomes, flowers, fruits, and seeds) being fairly uniform. Indeed, the infraspecific taxa recognized represent only part of the overall variation in the species, and one is faced with either recognizing poorly defined infraspecific taxa or treating the entire complex as a single polymorphic species. In the absence of thorough biosystematic and molecular studies on this group, we prefer not to recognize any infraspecific taxa. (Discussion copyrighted by Flora of North America; reprinted with permission.) |
Source | FNA vol. 7, p. 474. | FNA vol. 7, p. 469. |
Parent taxa | Brassicaceae > tribe Cardamineae > Cardamine | Brassicaceae > tribe Cardamineae > Cardamine |
Sibling taxa | ||
Synonyms | C. flexuosa subsp. debilis, C. flexuosa var. debilis, C. hirsuta subsp. flexuosa, C. scutata subsp. flexuosa | Dentaria californica, C. californica var. brevistyla, C. californica var. cardiophylla, C. californica subsp. cuneata, C. californica var. cuneata, C. californica var. fecunda, C. californica var. integrifolia, C. californica var. pubescens, C. californica var. robinsoniana, C. californica var. sinuata, C. cardiophylla, C. cuneata, C. integrifolia, C. integrifolia var. sinuata, C. pachystigma var. dissectifolia, C. paucisecta, C. sinuata, Dentaria californica var. cardiophylla, Dentaria californica var. cuneata, Dentaria californica var. integrifolia, Dentaria californica var. sinuata, Dentaria cardiophylla, Dentaria cuneata, Dentaria integrifolia, Dentaria integrifolia var. californica, Dentaria integrifolia var. cardiophylla, Dentaria integrifolia var. traceyi, Dentaria pachystigma var. dissectifolia, Dentaria sinuata |
Name authority | Withering: Arr. Brit. Pl. ed. 3, 3: 578. (1796) | (Nuttall) Greene: Fl. Francisc., 266. (1891) |
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