Flora of North America species comparison

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Cardamine

Cardamine holmgrenii

bitter-cress, toothwort

Holmgren's bitter-cress
Habit Annuals, biennials, or perennials; (perennials usually rhizomatous, sometimes tuberiform, stolons present in C. flagellifera, caudex present in C. bellidifolia); not scapose (subscapose in C. bellidifolia); glabrous or pubescent. Perennials; glabrous throughout.
Rhizomes

slender, 0.5–1 mm diam.
Stems

erect, ascending, decumbent, or prostrate, unbranched or branched.

erect to ascending, branched, 1.5–2 dm.
Leaves

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).

Cauline leaves

3 or 4, 3- or 5-foliolate, petiolate, leaflets petiolulate;

petiole 0.5–1.7 cm, base not auriculate;

lateral leaflet blade linear to linear-lanceolate, 0.9–1.5 cm × 1–3 mm, margins entire;

terminal leaflet (petiolule 0.3–0.5 cm), blade obovate to lanceolate (linear distally), 1.3–2.3 cm × 2–8 mm, base cuneate, margins entire or subapically 1- or 2-toothed.
Racemes

(corymbose or paniculate, bracteate in C. pattersonii), elongated in fruit.

ebracteate.
Flowers

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 oblong, 1.2–1.5 × 0.6–0.8 mm, lateral pair not saccate basally;

petals white, oblanceolate, 2–2.5 × 0.8–1 mm, (not clawed, apex rounded);

filaments: median pairs 1.5–1.8 mm, lateral pair 1–1.2 mm;

anthers ovate, 0.2–0.3 mm.
Fruiting pedicels

erect, ascending, divaricate, or reflexed, slender or stout.

ascending to suberect, 5–14 mm.
Fruits

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, 1–2 cm × ca. 1 mm;

ovules 16–24 per ovary;

style 0.5–0.7 mm.
Seeds

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.

brown, oblong, 1.2–1.5 × 0.8–1 mm.
Rhizomal

and basal leaves absent.
x

= 7, 8.

Cardamine

Cardamine holmgrenii
Phenology Flowering Jul–Aug.
Habitat Boggy slopes
Distribution
from USDA
Nearly worldwide
[BONAP county map]
from FNA
OR
[BONAP county map]
Discussion

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.)

Cardamine holmgrenii is known only from the type collection, from the Blue Mountains in Baker County.

(Discussion copyrighted by Flora of North America; reprinted with permission.)
Key
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
Source FNA vol. 7, p. 464. Authors: Ihsan A. Al-Shehbaz, Karol Marhold, Judita Lihová. FNA vol. 7, p. 475.
Parent taxa Brassicaceae > tribe Cardamineae Brassicaceae > tribe Cardamineae > Cardamine
Sibling taxa
C. angulata, C. angustata, C. bellidifolia, C. blaisdellii, C. breweri, C. bulbosa, C. californica, C. clematitis, C. concatenata, C. constancei, C. cordifolia, C. digitata, C. diphylla, C. dissecta, C. douglassii, C. flagellifera, C. flexuosa, C. hirsuta, C. impatiens, C. longii, C. macrocarpa, C. maxima, C. micranthera, C. microphylla, C. nuttallii, C. nymanii, C. occidentalis, C. oligosperma, C. pachystigma, C. parviflora, C. pattersonii, C. penduliflora, C. pensylvanica, C. pratensis, C. purpurea, C. rotundifolia, C. rupicola, C. umbellata
Subordinate taxa
C. angulata, C. angustata, C. bellidifolia, C. blaisdellii, C. breweri, C. bulbosa, C. californica, C. clematitis, C. concatenata, C. constancei, C. cordifolia, C. digitata, C. diphylla, C. dissecta, C. douglassii, C. flagellifera, C. flexuosa, C. hirsuta, C. holmgrenii, C. impatiens, C. longii, C. macrocarpa, C. maxima, C. micranthera, C. microphylla, C. nuttallii, C. nymanii, C. occidentalis, C. oligosperma, C. pachystigma, C. parviflora, C. pattersonii, C. penduliflora, C. pensylvanica, C. pratensis, C. purpurea, C. rotundifolia, C. rupicola, C. umbellata
Synonyms Dentaria, Dracamine, Loxostemon
Name authority Linnaeus: Sp. Pl. 2: 654. (1753): Gen. Pl. ed. 5, 295. (1754) Al-Shehbaz: Harvard Pap. Bot. 11: 275. (2007)
Web links 
Unless otherwise noted, all data comes from the online version of the Flora of North America, who have made most of their content available via a Creative Commons license (CC BY 4.0).



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