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large toothwort

California toothwort, coast toothwort, milk maids, tooth wort

Habit Perennials; glabrous (except leaflet margins and, sometimes, rachis). Perennials; usually glabrous, rarely minutely pubescent.
Rhizomes

cylindrical, 3–6 mm diam., (distinctly constricted at intervals, non-uniform diam., fleshy, slightly fragile, with dentate leaf scars).

(tuberiform, fragile), globose to ovoid or suboblong, (3–)4–10 mm diam., (fleshy, deeply underground).

Stems

erect, unbranched, 0.9–3(–4) dm.

erect, unbranched, (2–)2.7–6(–7)dm, usually glabrous, rarely pubescent.

Rhizomal leaves

3-foliolate, 7–20 cm, leaflets petiolulate or subsessile;

petiole 4–15 cm;

lateral leaflets subsessile or petiolulate, blade often similar to terminal, base often oblique;

terminal leaflet (petiolule 0.2–1(–1.7) cm), blade broadly ovate to oblong, 2–7.5 cm × 12–37 mm, base cuneate to obtuse, margins coarsely dentate to sharply incised, or deeply cleft into 2 or 3 lobes (lobes dentate or incised, margins puberulent).

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).

Cauline leaves

2 or 3, 3-foliolate, (rarely subopposite), petiolate, leaflets petiolulate or subsessile;

petiole (0.5–)1–4(–6.5) cm, base not auriculate;

lateral and terminal leaflets similar to rhizomal, distalmost sometimes much smaller.

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 (erect to ascending), oblong, 5–7 × 2–3 mm, lateral pair slightly saccate basally;

petals white or pink, oblanceolate, 10–17 × 3–6 mm, (not clawed, apex rounded);

filaments: median pairs 4–8 mm, lateral pair 3–6.5 mm;

anthers linear, 1.7–2.7 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

(flowering ones) horizontal to divaricate or deflexed, 7–20 mm.

ascending to divaricate, 10–33(–41) mm.

Fruits

(undeveloped), linear-lanceolate, to 3 cm × 2 mm;

ovules 10–14 per ovary;

style 3.5–7 mm.

linear, 2.2–5.4(–6) cm × 2–3 mm;

ovules 12–22 per ovary;

style 2–5(–6) mm.

Seeds

not known.

dark brown, oblong to broadly ovoid, 1.7–2.8 × 1.2–1.8 mm.

2n

= 120, 124, 132, 138, 156, 161, ca. 208.

= 32.

Cardamine maxima

Cardamine californica

Phenology Flowering Apr–Jun. Flowering Jan–May.
Habitat Rich woods, shady ravines, ledges, moist alluvial bottoms, steep forested slopes, stream banks Wooded ravines, forest floors, shady slopes, open woods, shady rock crevices, stream banks and bottoms, canyons, moist hillsides, cliffs
Elevation 0-1400 m (0-4600 ft)
Distribution
from FNA
CT; MA; ME; MI; NJ; NY; OH; PA; VT; NB; ON; QC
[WildflowerSearch map]
[BONAP county map]
from FNA
CA; Mexico (Baja California)
[WildflowerSearch map]
[BONAP county map]
Discussion

Cardamine maxima has not been found with mature fruits and seeds and it has long been suspected to be a hybrid between C. concatenata and C. diphylla. Molecular studies (P. W. Sweeney and R. A. Price 2000) indicate that C. maxima is distinct from both those species. Although we hesitate to maintain it as a species, its wide distribution and morphological distinctness warrant its recognition.

(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. 477. FNA vol. 7, p. 469.
Parent taxa Brassicaceae > tribe Cardamineae > Cardamine 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. holmgrenii, C. impatiens, C. longii, C. macrocarpa, 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
C. angulata, C. angustata, C. bellidifolia, C. blaisdellii, C. breweri, C. bulbosa, 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 maxima, C. anomala, Dentaria anomala 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 (Nuttall) Alph. Wood: Amer. Bot. Fl., 38. (1870) (Nuttall) Greene: Fl. Francisc., 266. (1891)
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