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FNA | Family List | FNA Vol. 7 | Brassicaceae

40. Cardamine Linnaeus, Sp. Pl. 2: 654. 1753; Gen. Pl. ed. 5, 295. 1754.

Bittercress [Greek kardamon, name for a cress] Bittercress [Greek kardamon, name for a cress]

Ihsan A. Al-Shehbaz, Karol Marhold, Judita Lihová

Dentaria Linnaeus; Dracamine Nieuwland; Loxostemon Hooker f. & Thomson

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. Stems erect, ascending, decumbent, or prostrate, unbranched or branched. 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). Racemes (corymbose or paniculate, bracteate in C. pattersonii), elongated in fruit. Fruiting pedicels erect, ascending, divaricate, or reflexed, slender or stout. 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. 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. 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. x = 7, 8.

Species ca. 200 (39 in the flora): nearly worldwide.

SELECTED REFERENCES Detling, L. E. 1936. The genus Dentaria in the Pacific states. Amer. J. Bot. 23: 570-576. Detling, L. E. 1937. The Pacific coast species of Cardamine. Amer. J. Bot. 24: 70-76. Franzke, A. and H. Hurka. 2000. Molecular systematics and biogeography of the Cardamine pratensis complex (Brassicaceae). Pl. Syst. Evol. 224: 213-234. Franzke, A. et al. 1998. Molecular systematics of Cardamine and allied genera (Brassicaceae): ITS and non-coding chloroplast DNA. Folia Geobot. 33: 225-240. Harriman, N. A. 1965. The Genus Dentaria L. (Cruciferae) in Eastern North America. Ph.D. dissertation. Vanderbilt University. Hart, T. W. and W. H. Eshbaugh. 1976. The biosystematics of Cardamine bulbosa (Muhl.) B.S.P. and C. douglasii Britt. Rhodora 78: 329-419. Lihová, J. and K. Marhold. 2006. Phylogenetic and diversity patterns in Cardamine (Brassicaceae)—A genus with conspicuous polyploid and reticulate evolution. In: A. K. Sharma and A. Sharma, eds. 2003+. Plant Genome: Biodiversity and Evolution. 2+ vols. in parts. Enfield, N.H. Vol. 1, part C, pp. 149-186. Lihová, J., K. Marhold, H. Kudoh, and M. Koch. 2006. Worldwide phylogeny and biogeography of Cardamine flexuosa (Brassicaceae) and its relatives. Amer. J. Bot. 93: 1206-1221. Lövkvist, B. 1956. The Cardamine pratensis complex—Outlines of its cytogenetics and taxonomy. Symb. Bot. Upsal. 14(2): 1-131. Montgomery, F. H. 1955. Preliminary studies in the genus Dentaria in eastern North America. Rhodora 57: 161-173. Schulz, O. E. 1903. Monographie der Gattung Cardamine. Bot. Jahrb. Syst. 32: 280-623. Sweeney, P. W. and R. A. Price. 2000. Polyphyly of the genus Dentaria (Brassicaceae): Evidence from trnL intron and ndhF sequence data. Syst. Bot. 25: 468-478.


1 Racemes bracteate.   32 Cardamine pattersonii
+ Racemes ebracteate   (2)
       
2 (1) Cauline leaves (at least some) simple (rarely absent)   (3)
+ Cauline leaves pinnately or palmately compound, 3-25-foliolate, sometimes pinnatisect and appearing compound   (14)
       
3 (2) Petioles of cauline leaves auriculate basally.   8 Cardamine clematitis (in part)
+ Petioles of cauline leaves not auriculate basally (rarely these leaves absent)   (4)
       
4 (3) Plants with distinct caudices; rhizomes absent; cauline leaves 0 or 1 (or 2).   3 Cardamine bellidifolia
+ Plants without caudices; rhizomes present; cauline leaves 2-23 (sometimes 1 in C. purpurea)   (5)
       
5 (4) Petals 0-0.7 mm; fruiting pedicels 0.5-2(-4) mm.   21 Cardamine longii
+ Petals 3.5-28 mm; fruiting pedicels (5-)7-50 mm   (6)
       
6 (5) Ovules 40-80 per ovary.   37 Cardamine rotundifolia
+ Ovules 10-24 per ovary   (7)
       
7 (6) 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.   24 Cardamine micranthera
+ 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 (7) Rhizomes (not fleshy), 1-3 mm diam.; rhizomal leaves usually absent   (9)
+ Rhizomes (fleshy), (3-)4-18 mm diam.; rhizomal leaves present   (11)
       
9 (8) Petals 15-28 mm; sepals 6-8 mm.   10 Cardamine constancei
+ Petals 5-12 mm; sepals 2-4.5 mm   (10)
       
10 (9) 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.   11 Cardamine cordifolia
+ 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.   36 Cardamine purpurea (in part)
       
11 (8) Petals 3-5 mm wide; fruits 1.4-2 mm wide; e, c North America   (12)
+ Petals 4-8 mm wide; fruits 2-4 mm; California   (13)
       
12 (11) Petals usually white, rarely pale pink; stem distally glabrous or trichomes 0.02-0.1 mm.   6 Cardamine bulbosa
+ Petals usually purple or pink, rarely white; stem distally with trichomes (0.2-)0.3-0.6(-0.8) mm.   15 Cardamine douglassii
       
13 (11) 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.   7 Cardamine californica (in part)
+ Petals usually purple or pink, rarely white, 14-18 mm; ovules 10-14 per ovary; stems 1-3 dm; rhizomal leaves simple.   30 Cardamine pachystigma
       
14 (2) Petiole of cauline leaves basally auriculate   (15)
+ Petiole of cauline leaves not basally auriculate   (16)
       
15 (14) Cauline leaves 3-foliolate; perennials (rhizomatous).   8 Cardamine clematitis (in part)
+ Cauline leaves (9-)13-25-foliolate; annuals or biennials.   20 Cardamine impatiens
       
16 (14) Annuals or biennials, not rhizomatous   (17)
+ Perennials, rhizomatous   (22)
       
17 (16) Rachises strongly flexuous to geniculate; petals 6-8 mm; fruits (2.5-)3-4.6 × 1.7-2.1 mm.   22 Cardamine macrocarpa
+ 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 (17) Basal leaves rosulate, persistent to anthesis   (19)
+ Basal leaves not rosulate, often withered by anthesis   (20)
       
19 (18) Stamens usually 4, rarely 5 or 6; fruiting pedicels erect to ascending; fruitsoften appressed to rachis, valves glabrous; seeds narrowly margined.   18 Cardamine hirsuta
+ Stamens 6; fruiting pedicels divaricate-ascending; fruits often not appressed to rachis, valves sparsely pubescent or glabrous; seeds often not margined (wingless).   29 Cardamine oligosperma
       
20 (18) Lateral leaflets of cauline leaves narrowly oblong, linear or filiform, 1-3 mm wide; fruits 0.6-0.9 mm wide.   31 Cardamine parviflora
+ Lateral leaflets of cauline leaves orbicular, ovate, elliptic, or oblong, 4-25 mm wide; fruits 0.8-1.5 mm wide   (21)
       
21 (20) Rachises slightly flexuous; ovules 18-40 per ovary; plants of waste grounds, disturbed sites.   17 Cardamine flexuosa
+ Rachises straight; ovules 40-80 per ovary; plants of margins of wet habitats.   34 Cardamine pensylvanica
       
22 (16) Rhizomes tuberiform, usually segmented, fragile (segments fusiform, ovoid, oblong, or globose)   (23)
+ Rhizomes cylindrical, segments not fragile   (29)
       
23 (22) Cauline leaves biternate, distal lobes linear.   14 Cardamine dissecta
+ Cauline leaves 3-11-foliolate, distal lobes not linear   (24)
       
24 (23) Cauline leaves 5-11-foliolate.   33 Cardamine penduliflora
+ Cauline leaves usually 3-, sometimes 5-7-foliolate, rarely simple   (25)
       
25 (24) Rhizomes moniliform; e, c United States   (26)
+ Rhizomes not moniliform; Pacific North America   (27)
       
26 (25) Cauline leaves different from rhizomal leaves, usually alternate, rarely opposite.   2 Cardamine angustata
+ Cauline leaves similar to rhizomal leaves, usually whorled or opposite, rarely alternate.   9 Cardamine concatenata
       
27 (25) Middle cauline leaves 5- or 7-foliolate; sepals 1.7-2 mm; petals 4-6 mm.   28 Cardamine occidentalis
+ Middle cauline leaves 3(-5)-foliolate; sepals 3.5-5.5 mm; petals 8-15 mm   (28)
       
28 (27) Cauline leaves 2-5; stems (2-)2.7-6(-7) dm; ovules 12-22 per ovary; petals white or pale rose; California.   7 Cardamine californica (in part)
+ 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.   26 Cardamine nuttallii
       
29 (22) Cauline leaves pinnatisect or (7-)9-21-foliolate   (30)
+ Cauline leaves 3-5(-7)-foliolate   (31)
       
30 (29) Leaves thick, veins impressed.   27 Cardamine nymanii
+ Leaves thin, veins raised.   35 Cardamine pratensis
       
31 (29) Rhizomes compact at stem base, stolons present.   16 Cardamine flagellifera
+ Rhizomes slender, stolons absent   (32)
       
32 (31) Rhizomes fleshy, with dentate leaf scars; plants of North America east of the 110th meridian   (33)
+ Rhizomes not fleshy, without dentate leaf scars; plants of North America west of the 110th meridian   (34)
       
33 (32) Rhizomes somewhat uniform in diameter; cauline leaves (sub) opposite.   13 Cardamine diphylla
+ Rhizomes not uniform in diameter (distinctly constricted at intervals); cauline leaves usually alternate, rarely subopposite.   23 Cardamine maxima
       
34 (32) Plants cespitose, hirsute; petals purple or pink, rarely white.   36 Cardamine purpurea (in part)
+ Plants not cespitose, glabrous or sparsely pubescent or pilose; petals usually white, rarely pink   (35)
       
35 (34) Basal leaves rosulate; fruiting racemes subumbellate.   39 Cardamine umbellata
+ Basal leaves absent; fruiting racemes not subumbellate   (36)
       
36 (35) Sepals 1.2-1.5 mm; petals 2-2.5 × 0.8-1 mm; anthers 0.2-0.3 mm.   19 Cardamine holmgrenii
+ Sepals 2-5 mm; petals 3.5-15 × 1.5-8 mm; anthers 0.7-1.5 mm   (37)
       
37 (36) Rhizomal leaves absent; ovules 14-28 per ovary.   5 Cardamine breweri
+ Rhizomal leaves present; ovules 6-16(-24) per ovary   (38)
       
38 (37) Stems hirsute at base; cauline leaves (3 or) 4-8.   1 Cardamine angulata
+ Stems usually glabrous at base, rarely sparsely pilose; cauline leaves 1-3 (or 4)   (39)
       
39 (38) Rhizomal leaves palmately or subpalmately compound, fleshy; Montana.   38 Cardamine rupicola
+ Rhizomal leaves pinnately compound, not fleshy; Alaska, Northwest Territories, Yukon   (40)
       
40 (39) Rhizomal leaves with terminal leaflet blade broadly obovateto suborbicular, margins often 3-5-toothed.   4 Cardamine blaisdellii
+ Rhizomal leaves with terminal leaflet blade linear, narrowly oblong, narrowly oblanceolate, ovate, or elliptic, margins entire   (41)
       
41 (40) Cauline leaves with terminal leaflet narrowly lanceolate to linear, 1-3.2(-4) cm.   12 Cardamine digitata
+ Cauline leaves with terminal leaflet ovate to elliptic, 0.6-1.3 cm.   25 Cardamine microphylla

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