Cardamine
Cardamine pratensis
bitter-cress, toothwort
Brewer's bitter-cress, cuckoo bitter-cress, cuckoo flower, European field bitter-cress, lady's-smock, meadow bittercress, pink cuckoo bitter-cress
cylindrical, (not fragile), relatively short, (not fleshy).
erect, ascending, decumbent, or prostrate, unbranched or branched.
erect, unbranched, (0.8–)1.5–5.5(–8) dm.
cauline, rhizomal, or basal;
rhizomal and basal rosulate or not, petiolate, blade margins entire, toothed, or 1–3-pinnatisect, or palmately lobed, sometimes trifoliolate, pinnately, palmately, or bipinnately compound (leaflets petiolulate, subsessile, or sessile);
cauline (usually alternate, rarely opposite or whorled) petiolate or sessile, blade (base cuneate, attenuate, or auriculate to sagittate), margins entire, dentate, or variously lobed, (leaflets petiolulate or sessile).
simple or 5–9(–31)-foliolate, to 30 cm, (thin, veins raised), leaflets petiolulate or sessile;
petiole (1–)2–7(–10) cm;
lateral leaflets petiolulate or sessile, blade similar to terminal or smaller, orbicular, ovate, or obovate, margins crenate or repand;
terminal leaflet (petiolule to 1.5 cm), blade orbicular or broadly obovate, 0.3–2 cm diam., base usually rounded, rarely subreniform or cuneate, margins repand, (apex rounded).
2–12(–18), pinnatisect, petiolate, leaflets petiolulate or sessile, (2–17 cm including petiole, thin, veins raised);
petiole base not auriculate;
lobes or leaflets (4–7(–13) each side of rachis), petiolulate or sessile and decurrent, blade similar to terminal lobe or leaflet, margins usually entire, rarely dentate;
terminal lobe or leaflet (petiolulate or sessile), blade linear, oblong, ovate, or lanceolate, 1–2.5(–3.5) cm × 5–8(–10) mm, (surfaces glabrous).
(corymbose or paniculate, bracteate in C. pattersonii), elongated in fruit.
ebracteate.
sepals (caducous), usually erect, rarely spreading or ascending, ovate or oblong, lateral pair saccate or not basally, (usually glabrous, rarely pubescent);
petals (rarely absent), white, pink, purple, or lilac, obovate, spatulate, or oblanceolate, claw absent or strongly differentiated from blade, (apex obtuse, rounded, emarginate, or subemarginate);
stamens (6, rarely 4), equal in length;
filaments not dilated basally;
anthers ovate, oblong, or linear, (apex obtuse), glabrous [rarely pubescent];
nectar glands confluent, lateral glands annular or semiannular, subtending bases of stamens, median glands present (2, rarely 4) or absent.
sepals (erect or spreading), oblong or ovate, (2.5–)3–5(–6) × 1–2 mm, lateral pair saccate basally;
petals usually purple or lilac, rarely white, obovate, (6–)8–15(–18) × 3–7.5(–10) mm, (clawed, apex rounded or emarginate);
filaments: median pairs 5–10 mm, lateral pair 3–6 mm;
anthers narrowly oblong, (0.8–)1.2–2 mm.
erect, ascending, divaricate, or reflexed, slender or stout.
erect-ascending or subdivaricate, (5–)12–25(–30) mm.
siliques, sessile, usually linear, rarely narrowly oblong or narrowly lanceolate, smooth or torulose, latiseptate;
valves (papery, elastically dehiscent, becoming spirally or circinately coiled) each not veined, glabrous or, rarely, pubescent;
replum strongly flattened;
septum complete, (membranous);
ovules 4–80 per ovary;
style usually distinct, rarely obsolete;
stigma capitate.
linear, (1.6–)2.5–4.5(–5) cm × (1.2–)1.5–2.3 mm;
ovules 20–30 per ovary;
style (0.5–)1–2.2(–2.7) mm, (stout).
uniseriate, flattened, usually not winged, rarely margined or winged, oblong, ovoid, or globose;
seed coat (smooth, minutely reticulate, colliculate, or rugose) mucilaginous or not when wetted;
cotyledons accumbent, rarely incumbent.
light brown, oblong, 1.2–1.8(–2) × 1–1.4 mm.
= 7, 8.
= 16.
Cardamine
Cardamine pratensis
Species ca. 200 (39 in the flora).
R. C. Rollins (1993) recognized 50 species and 18 varieties of Cardamine in North America minus Greenland. Of these, 14 species are restricted to Central America and one, C. ×incisa (Eames) K. Schumann, is not recognized here. The oversplitting of some species into poorly delimited varieties is avoided in this account, and four species (C. blaisdellii, C. holmgrenii, C. nymanii, C. umbellata) are added to the flora. Cardamine corymbosa Hooker f., a native of New Zealand, apparently is becoming naturalized in greenhouses and in some gardens in British Columbia, as evidenced from Lomer 4368 (MO), a specimen collected in Sechelt Peninsula on 6 June 2002.
(Discussion copyrighted by Flora of North America; reprinted with permission.)
The taxonomy of Cardamine pratensis in North America requires further detailed study. Most, if not all, populations of this species were introduced from Europe. Some specimens resemble the European C. dentata Schultes (high polyploid, characterized by all leaves, including distalmost, pinnate with petiolate and sometimes deciduous leaflets) and these populations might be native.
(Discussion copyrighted by Flora of North America; reprinted with permission.)
1. Racemes bracteate. | C. pattersonii |
1. Racemes ebracteate | → 2 |
2. Cauline leaves (at least some) simple (rarely absent) | → 3 |
2. Cauline leaves pinnately or palmately compound, 3-25-foliolate, sometimes pinnatisect and appearing compound | → 14 |
3. Petioles of cauline leaves auriculate basally. | C. clematitis |
3. Petioles of cauline leaves not auriculate basally (rarely these leaves absent) | → 4 |
4. Plants with distinct caudices; rhizomes absent; cauline leaves 0 or 1 (or 2). | C. bellidifolia |
4. Plants without caudices; rhizomes present; cauline leaves 2-23 (sometimes 1 in C. purpurea) | → 5 |
5. Petals 0-0.7 mm; fruiting pedicels 0.5-2(-4) mm. | C. longii |
5. Petals 3.5-28 mm; fruiting pedicels (5-)7-50 mm | → 6 |
6. Ovules 40-80 per ovary. | C. rotundifolia |
6. Ovules 10-24 per ovary | → 7 |
7. Fruits 0.8-1.6 cm × 0.8-1 mm; anthers ovate, ca. 0.2 mm; petals 3.5-5 × 1.2-1.8 mm. | C. micranthera |
7. Fruits (1.5-)2-6 cm × 1.2-4 mm; anthers oblong to linear, 0.6-3 mm; petals (5-)7-28 × 3-7 mm | → 8 |
8. Rhizomes (not fleshy), 1-3 mm diam.; rhizomal leaves usually absent | → 9 |
8. Rhizomes (fleshy), (3-)4-18 mm diam.; rhizomal leaves present | → 11 |
9. Petals 15-28 mm; sepals 6-8 mm. | C. constancei |
9. Petals 5-12 mm; sepals 2-4.5 mm | → 10 |
10. Cauline leaves (3-)5-17(-23); petals white, 7-12 mm; fruits (2-)2.5-3.7(-4) cm; stems 2-7(-10.2) dm. | C. cordifolia |
10. Cauline leaves 1-3; petals usually purple to pink, rarely white, 5-7(-9) mm; fruits 1.5-2.5 cm; stems to 1.5 dm. | C. purpurea |
11. Petals 3-5 mm wide; fruits 1.4-2 mm wide; e, c North America | → 12 |
11. Petals 4-8 mm wide; fruits 2-4 mm; California | → 13 |
12. Petals usually white, rarely pale pink; stem distally glabrous or trichomes 0.02-0.1 mm. | C. bulbosa |
12. Petals usually purple or pink, rarely white; stem distally with trichomes (0.2-)0.3-0.6(-0.8) mm. | C. douglassii |
13. Petals white or pale rose, rarely purple, 8-13(-17) mm; ovules 12-22 per ovary; stems (2-)2.7-6(-7) dm; rhizomal leaves usually 3(-7)-foliolate, rarely simple. | C. californica |
13. Petals usually purple or pink, rarely white, 14-18 mm; ovules 10-14 per ovary; stems 1-3 dm; rhizomal leaves simple. | C. pachystigma |
14. Petiole of cauline leaves basally auriculate | → 15 |
14. Petiole of cauline leaves not basally auriculate | → 16 |
15. Cauline leaves 3-foliolate; perennials (rhizomatous). | C. clematitis |
15. Cauline leaves (9-)13-25-foliolate; annuals or biennials. | C. impatiens |
16. Annuals or biennials, not rhizomatous | → 17 |
16. Perennials, rhizomatous | → 22 |
17. Rachises strongly flexuous to geniculate; petals 6-8 mm; fruits (2.5-)3-4.6 × 1.7-2.1 mm. | C. macrocarpa |
17. Rachises not flexuous (slightly so in C. flexuosa); petals 1.5-4(-5) mm; fruits (0.5-) 1-2.5(-3.2) × 0.6-1.5 mm | → 18 |
18. Basal leaves rosulate, persistent to anthesis | → 19 |
18. Basal leaves not rosulate, often withered by anthesis | → 20 |
19. Stamens usually 4, rarely 5 or 6; fruiting pedicels erect to ascending; fruitsoften appressed to rachis, valves glabrous; seeds narrowly margined. | C. hirsuta |
19. Stamens 6; fruiting pedicels divaricate-ascending; fruits often not appressed to rachis, valves sparsely pubescent or glabrous; seeds often not margined (wingless). | C. oligosperma |
20. Lateral leaflets of cauline leaves narrowly oblong, linear or filiform, 1-3 mm wide; fruits 0.6-0.9 mm wide. | C. parviflora |
20. Lateral leaflets of cauline leaves orbicular, ovate, elliptic, or oblong, 4-25 mm wide; fruits 0.8-1.5 mm wide | → 21 |
21. Rachises slightly flexuous; ovules 18-40 per ovary; plants of waste grounds, disturbed sites. | C. flexuosa |
21. Rachises straight; ovules 40-80 per ovary; plants of margins of wet habitats. | C. pensylvanica |
22. Rhizomes tuberiform, usually segmented, fragile (segments fusiform, ovoid, oblong, or globose) | → 23 |
22. Rhizomes cylindrical, segments not fragile | → 29 |
23. Cauline leaves biternate, distal lobes linear. | C. dissecta |
23. Cauline leaves 3-11-foliolate, distal lobes not linear | → 24 |
24. Cauline leaves 5-11-foliolate. | C. penduliflora |
24. Cauline leaves usually 3-, sometimes 5-7-foliolate, rarely simple | → 25 |
25. Rhizomes moniliform; e, c United States | → 26 |
25. Rhizomes not moniliform; Pacific North America | → 27 |
26. Cauline leaves different from rhizomal leaves, usually alternate, rarely opposite. | C. angustata |
26. Cauline leaves similar to rhizomal leaves, usually whorled or opposite, rarely alternate. | C. concatenata |
27. Middle cauline leaves 5- or 7-foliolate; sepals 1.7-2 mm; petals 4-6 mm. | C. occidentalis |
27. Middle cauline leaves 3(-5)-foliolate; sepals 3.5-5.5 mm; petals 8-15 mm | → 28 |
28. Cauline leaves 2-5; stems (2-)2.7-6(-7) dm; ovules 12-22 per ovary; petals white or pale rose; California. | C. californica |
28. Cauline leaves 1-3; stems 0.5-2(-3) dm; ovules 8-16 per ovary; petals usually purple to pale pink, rarely white; British Columbia, California, Oregon, Washington. | C. nuttallii |
29. Cauline leaves pinnatisect or (7-)9-21-foliolate | → 30 |
29. Cauline leaves 3-5(-7)-foliolate | → 31 |
30. Leaves thick, veins impressed. | C. nymanii |
30. Leaves thin, veins raised. | C. pratensis |
31. Rhizomes compact at stem base, stolons present. | C. flagellifera |
31. Rhizomes slender, stolons absent | → 32 |
32. Rhizomes fleshy, with dentate leaf scars; plants of North America east of the 110th meridian | → 33 |
32. Rhizomes not fleshy, without dentate leaf scars; plants of North America west of the 110th meridian | → 34 |
33. Rhizomes somewhat uniform in diameter; cauline leaves (sub) opposite. | C. diphylla |
33. Rhizomes not uniform in diameter (distinctly constricted at intervals); cauline leaves usually alternate, rarely subopposite. | C. maxima |
34. Plants cespitose, hirsute; petals purple or pink, rarely white. | C. purpurea |
34. Plants not cespitose, glabrous or sparsely pubescent or pilose; petals usually white, rarely pink | → 35 |
35. Basal leaves rosulate; fruiting racemes subumbellate. | C. umbellata |
35. Basal leaves absent; fruiting racemes not subumbellate | → 36 |
36. Sepals 1.2-1.5 mm; petals 2-2.5 × 0.8-1 mm; anthers 0.2-0.3 mm. | C. holmgrenii |
36. Sepals 2-5 mm; petals 3.5-15 × 1.5-8 mm; anthers 0.7-1.5 mm | → 37 |
37. Rhizomal leaves absent; ovules 14-28 per ovary. | C. breweri |
37. Rhizomal leaves present; ovules 6-16(-24) per ovary | → 38 |
38. Stems hirsute at base; cauline leaves (3 or) 4-8. | C. angulata |
38. Stems usually glabrous at base, rarely sparsely pilose; cauline leaves 1-3 (or 4) | → 39 |
39. Rhizomal leaves palmately or subpalmately compound, fleshy; Montana. | C. rupicola |
39. Rhizomal leaves pinnately compound, not fleshy; Alaska, Northwest Territories, Yukon | → 40 |
40. Rhizomal leaves with terminal leaflet blade broadly obovateto suborbicular, margins often 3-5-toothed. | C. blaisdellii |
40. Rhizomal leaves with terminal leaflet blade linear, narrowly oblong, narrowly oblanceolate, ovate, or elliptic, margins entire | → 41 |
41. Cauline leaves with terminal leaflet narrowly lanceolate to linear, 1-3.2(-4) cm. | C. digitata |
41. Cauline leaves with terminal leaflet ovate to elliptic, 0.6-1.3 cm. | C. microphylla |
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