Classification
 Nomenclature
Scientific Name:
Cardamine L., Sp. Pl. 654 (1753)
Synonymy:
  • = Iti Garn.-Jones & P.N.Johnson, New Zealand J. Bot. 25: 603 (1987)
 Description

Herbs annual or perennial. Trichomes absent or present, when present simple. Stems erect, ascending, or decumbent, sometimes rhizomatous, branched or unbranched. Basal leaves rosulate, simple, pinnatifid or pinnatisect, petiolate; lateral leaflets petiolulate, subsessile or sessile. Cauline leaves present or absent, when present alternate, simple, pinnatifid or pinnatisect as basal leaves but usually smaller and narrower. Inflorescences corymbose or racemose, flowers sometimes solitary; ebracteate, elongated in fruit. Fruiting pedicels long and slender or shorter and thickened, erect or spreading. Sepals usually saccate, margin usually membranous. Petals absent or present; when present usually white, sometimes pink or purple; claw usually differentiated from blade and longer than sepals. Filaments 6 and tetradynamous, sometimes 2 or 4 and then equal in length, linear or rarely dilated. Anthers usually yellow or sometimes maroon, glabrous, held at a similar height to stigma. Nectar glands confluent and subtending bases of all stamens. Ovules few to many. Style distinct or sometimes indistinct; stigma capitate, entire. Fruit explosively dehiscent silique or an indehiscent silicle; weakly latiseptate or ± terete, smooth or weakly torulose, sessile; valves not veined, glabrous or hairy, dehiscing elastically acropetally, coiled or indehiscent and remaining flattened; replum strongly flattened; septum complete, membranous, translucent. Seeds uniseriate, winged or wingless, flattened; seed coat minutely reticulate; embryo white, venation rarely violet-purple.

 Taxonomy

Cardamine belongs to the family Brassicaceae, tribe Cardamineae, a tribe of 10 genera and over 340 species (Al-Shehbaz et al. 2006). Cardamine and Rorippa are the largest genera of the tribe, with Cardamine comprising about 230 species (Al-Shehbaz et al. 2006; Heenan 2017).

Species of Cardamine in New Zealand are morphologically diverse and offer a variety of morphological characters for taxonomic circumscription and identification (Heenan 2017). Some of the more important taxonomic characters include: leaves simple or pinnate and coriaceous or membranous, the presence or absence of leaflet axillary hydathodes, the length and width of the sepals, petals, siliques and seeds, seed size and shape, and the presence or absence of a margin wing.

 Key
1Fruit a silicle or silique, <4.0 mm long2
Fruit a silique, >6.0 mm long3
2Leaves broadly orbicular, <2.2 mm long; 2 stamens, bent at distal end; silique <4.0 mm longcubita
Leaves simple and linear-spathulate or pinnatifid with narrow-spathulate lobes, 20.0–50.0 mm long; 4(–6) stamens, straight; silicle <3.5 mm longlacustris
3Inflorescence decumbent or prostrate, axillary rosettes present in nodespanatohea
Inflorescence erect or spreading, axillary rosettes absent from nodes4
4Leaves simple with a length to width ratio of >1.5 : 1, or pinnatifid with the terminal leaflet having a length to width ratio of >1.5 : 1 and 1–4 basal lobes, usually coriaceous5
Leaves simple with a length to width ratio ±1 : 1 or pinnatisect with the terminal leaflet having a length to width ratio ±1 : 1 and 1–14 lateral leaflets with distinct petiolules, usually membranous13
5Anthers maroon or yellow and flushed maroon6
Anthers yellow8
6Sepals 0.8–1.1 mm wide; petals 4.0–7.0 mm long; silique style 0.3–0.5 mm longdimidia
Sepals 1.3–1.8 mm wide; petals 7.0–11.0 mm long; silique style 1.0–3.5 mm long7
7Leaves red-brown to brown or occasionally green; filaments linear their entire length, 0.15–0.35 mm wide; silique style 1.7–3.5 mm longbilobata
Leaves grey-green to green; filaments conspicuously dilated at base, 0.35–0.65 mm wide; silique style 1.0–1.2 mm longdilatata
8Leaves always simple9
Leaves simple and pinnatisect or pinnatifid10
9Leaves membranous, matt, up to 90 mm long, lamina obovate, elliptic, broadly elliptic, petiole up to 60 mm long; sepals 1.9–2.0 mm long; petals 2.6–4.2 mm long; filaments 1.8–2.7 mm long; siliques 7.5–8.0 mm longalticola
Leaves coriaceous, glossy, up to 45 mm long, lamina narrowly obovate to spathulate, petiole up to 25 mm long; sepals 1.6–1.8 mm long; petals 2.1–3.2 mm long; filaments 1.2–1.3 mm long; siliques 7.0–14.0 mm longsciaphila
10Leaves usually pinnatisect or pinnatifid, sometimes simple, terminal lobe or simple pinna 2–8 mm long and 0.5–5.0 mm wide; cauline leaves occasionally present, usually pinnatifid or pinnatisect; inflorescence spreading to ascendingmutabilis
Leaves usually simple, sometimes pinnatifid, terminal lobe or simple pinna 3.0–45 mm long and 2.0–23.0 mm wide; cauline leaves usually simple; inflorescence ascending to erect11
11Leaves membranous, lamina surface glabrous or hairy; inflorescence usually short and held just above the leaves, flowers clustered towards distal end in a corymb; plants of Subantarctic Islandsdepressa
Leaves coriaceous, lamina surface glabrous; inflorescence usually elongated and held well above the leaves, flowers distributed along inflorescence, plants of South Island12
12Leaves grey-green, matt, 4.0–45.0 mm long, 2.5–23.0 mm wide; sepals 2.9–3.0 mm long; inflorescence up to 300 mm longpachyphylla
Leaves green, purple-brown, brown-green or bronze, glossy, 5.0–20.0 mm long, 2.0–9.5 mm wide; sepals 2.0–2.5 mm long; inflorescence up to 110 mm longdimidia
13Leaves with axillary hydathodes14
Leaves without axillary hydathodes20
14Siliques glabrous15
Siliques hairy18
15Leaves coriaceous, terminal leaflet 3.0–7.7 mm long and 2.8–8.6 mm wide; seed wing present, reticulate cells conspicuously thick-walledeminentia
Leaves membranous, terminal leaflet 3.5–21.0 mm long and 4.7–23.0 mm wide; seed wing absent, reticulate cells not obviously thick-walled16
16Lateral leaflets 1–2, green to dark green, with indistinct petiolules 0.2–1.5 mm long; petals broadly obovate, 4.2–4.5 mm wide; siliques 2.1–2.4 mm wide; seeds 1.0–1.2 mm widemegalantha
Lateral leaflets 2–8, light green to green, with distinct petiolules 2.0–20.0 mm long; petals obovate to narrowly obovate, 0.7–3.6 mm wide; siliques 0.7–1.4 mm wide; seeds 0.6–0.9 mm wide17
17Leaf axillary and marginal hydathodes usually associated with additional hairs; sepals 2.4–3.2 mm long; petals obovate, 4.5–8.0 mm long, 2.5–3.6 mm widecoronata
Leaf axillary and marginal hydathodes solitary and not associated with additional hairs; sepals 1.3–1.6 mm long; petals narrowly obovate, 2.8–4.5 mm long, 0.7–1.4 mm wideforsteri
18Terminal leaflet 2.6–7.0 mm long, 3.2–10.0 mm wide; lateral leaflets 2.0–5.0 mm long, 1.8–5.8 mm wide; leaflet margins entire to weakly lobed; siliques 13.0–25.0 mm longbisetosa
Terminal leaflet 3.5–21.0 mm long, 4.7–26.0 mm wide; lateral leaflets 2.0–20.0 mm long, 2.0–18.0 mm wide; leaflet margins shallowly to deeply lobed; siliques 20.0–38.0 mm long19
19Leaf axillary and marginal hydathodes usually associated with additional hairs; sepals 2.4–3.2 mm long; petals obovate, 4.5–8.0 mm long, 2.5–3.6 mm widecoronata
Leaf axillary and marginal hydathodes solitary and not associated with additional hairs; sepals 1.3–1.6 mm long; petals narrowly obovate, 2.8–4.5 mm long, 0.7–1.4 mm wideforsteri
20Seeds winged21
Seeds not winged23
21Leaves membranous; silique 1.5–2.3 mm wide; seed oblong to oblong-elliptic, green to green-brown, wing at apexunguiculus
Leaves coriaceous; silique 1.1–1.9 mm wide; seed orbicular, broadly oblong to oblong, henna, wing at apex and lateral margin22
22Leaves glossy, terminal pinna usually with prominent triangular lobes, petiolule <2.5 mm long; inflorescence up to 120 mm long; petals 2.6–7.5 mm long, 0.8–3.8 mm wide; seeds 0.9–1.5 mm longalalata
Leaves matt, terminal pinna usually irregularly lobed, petiolule 2.0–15.0 mm long; inflorescence up to 350 mm long; petals 5.0–11.0 mm long, 2.5–5.5 mm wide; seeds 1.2–2.1 mm longverna
23Siliques hairy24
Siliques glabrous28
24Silique hairs sparse, patent, 0.35–0.6 mm longhirsuta
Silique hairs moderate to dense, appressed to spreading, up to 0.35 mm long25
25Petals 4.5–8.0 mm long, 2.5–3.6 mm wide; sepals 2.4–3.2 mm long, 1.1–1.3 mm wide; median filaments 2.5–4.0 mm longcoronata
Petals 2.7–4.5 mm long, 0.7–1.4 mm wide; sepals 1.2–1.9 mm long, 0.5–0.9 mm wide; median filaments 1.6–2.7 mm long26
26Leaves 2.6–7.0 mm long, 3.2–10.0 mm wide, petiolule 0.5–5.3 mm long; petals 2.7–3.5 mm long, 0.7–0.8 mm wide; anthers cream to pale yellow or maroonbisetosa
Leaves 3.1–25.0 mm long, 4.7–25.0 mm wide, petiolule 2.0–16.0 mm long; petals 3.0–4.5 mm long, 0.7–1.4 mm wide; anthers cream or pale yellow27
27Sepals 1.7–1.9 mm long; median filaments 2.6–2.7 mm long, lateral filaments 2.2–2.3 mm long; silique 1.0–1.4 mm wide; seeds 1.0–1.2 mm longintonsa
Sepals 1.3–1.6 mm long; median filaments 1.6–2.5 mm long, lateral filaments 1.6–2.0 mm long; silique 0.7–1.1 mm long; seeds 1.2–1.4 mm longforsteri
28Inflorescence corymbose29
Inflorescence racemose40
29Petals >6.5 mm longreptans
Petals <6.5 mm long30
30Petals <3.5 mm long31
Petals 3.5–6.5 mm long33
31Leaves membranous, green or green-brown, terminal leaflet usually broader than long, adaxial surface usually sparsely to moderately hairy, margins with conspicuous patent hairs; sepals 0.4–0.6 mm wide, with conspicuous spreading hairs; siliques 0.7–0.8 mm wide, usually red-brown, weakly torulose, seed outlines usually visible through valve walls; seeds 0.5–0.7 mm wideheleniae
Leaves coriaceous or semi-coriaceous, green, terminal leaflet usually with ± equal length to width or longer than broad; adaxial surface glabrous, margin sometimes with a few sparse, inconspicuous hairs; sepals 0.5–1.0 mm wide, usually glabrous; siliques 0.9–1.5 mm wide, green, not torulose, seed outlines not visible through valve walls; seeds 0.9–1.0 mm wide32
32Leaves with conspicuous marginal hydathodes, terminal leaflet 3.0–11.0 mm long, 1.5–8.0 mm wide; siliques 5.7–14.0 mm long; style 0.3–0.5 mm longexigua
Leaves with inconspicuous marginal hydathodes, terminal leaflet 1.5–2.5 mm long, 1.0–2.5 mm wide; siliques 13.0–25.0 mm long; style 0.9–1.5 mm longparvula
33Terminal leaflet longer than wide, base attenuate to cuneate, occasionally obtuse34
Terminal leaflet similar length to width, base obtuse, truncate or cordate35
34Leaves usually simple, seldom pinnate, green, membranous, glabrous or sparsely hairy, with 0–2 lateral pinnae; terminal pinna oblong-elliptic, broadly oblong-elliptic to oblong-orbicular, base cuneate to occasionally obtuse; peduncle up to 150 mm long, glabrous; silique 9.0–20.0 mm longintegra
Leaves usually pinnate, seldom simple, green to green-brown, ± coriaceous, glabrous, with 0–6 lateral pinnae; terminal pinna elliptic, broadly elliptic, obovate, ovate or spathulate, base attenuate to cuneate; peduncle up to 30.0 mm long, glabrous or hairy; silique 6.0–11.0 mm longmutabilis
35Leaves coriaceus36
Leaves membranous37
36Leaves green-brown to purple-brown, lateral pinnae 0–2; terminal pinna 2.0–6.0 mm long, 1.8–8.0 mm wide; sepals 1.8–2.4 mm long; siliques 12.0–20.0 mm longserpentina
Leaves green to green-brown, lateral pinnae 0–4; terminal pinna 2.0–25.0 mm long, 2.0–30.0 mm wide; sepals 2.3–3.0 mm long; siliques 20.0–40.0 mm longglara
37Sepals 1.1–1.6 mm long; silique 8.0–13.5 mm longunicaulis
Sepals 1.3–3.0 mm long; silique 12.0–32.0 mm long38
38Inflorescence up to 180 mm long, with 1–9 corymbs, corymbs 2–19‑flowered, occasionally with solitary flowers; leaf terminal pinna 3.0–27.0 mm long, 3.0–30.0 mm widegrandiscapa
Inflorescence with 1–3 corymbs, corymbs 2–8-flowered, often with solitary flowers; leaf terminal pinna 2.0–18.0 mm long, 2.0–16.0 mm wide39
39Leaves light green, leaf margin usually shallowly sinuate; sepals 1.8–2.2 mm long; petals 2.1–3.0 mm wide; silique style 0.8–1.0 mm longsinuatifolia
Leaves green, leaf margin entire; sepals 1.4–2.0 mm long; petals 1.8–2.3 mm wide; silique style 0.2–0.4 mm longcorymbosa
40Lateral pinnae ≥1041
Lateral pinnae ≤843
41Terminal pinna 3.4–4.7 mm long, 3.5–4.1 mm wide; inflorescence up to 50 mm long; silique 1.7–2.2 mm widelatior
Terminal pinna 2–25 mm long, 2–27 mm wide; inflorescence up to 350 mm long; silique 1.0–1.7 mm wide42
42Leaf terminal pinna larger than lateral pinnae; sepals 2.5–4.0 mm long, 1.0–2.4 mm wide; petals 7.0–11.0 mm long, 2.1–6.0 mm wide; median filaments 3.2–4.0 mm long, lateral filaments 2.0–2.7 mm long; seeds 1.5–1.7 mm longpratensis
Leaf terminal pinna similar size to lateral pinnae; sepals 1.5–3.5 mm long, 0.7–1.4 mm wide; petals 2.5–4.0 mm long, 1.0–1.7 mm wide; median filaments 2.3–2.4 mm long, lateral filaments 2.7–2.7 mm long; seeds 0.8–1.0 mm longflexuosa
43Lateral pinnae ≤244
Lateral pinnae ≥346
44Leaves coriaceous, grey-green to blue-grey, broadly ovate or broadly orbicular-deltoid, base truncate; sepals sparsely hairycaesiella
Leaves membranous, green, pandurate, pandurate-orbicular or rhomboid-pandurate, base cordate; sepals glabrous45
45Sepals 2.7–3.1 mm long; petals 6.0–7.0 mm long, 4.2–4.5 mm wide; silique 2.1–2.4 mm wide; embryo cotyledons without coloured veinsmegalantha
Sepals 2.0–2.4 mm long; petals 4.1–4.5 mm long, 1.9–2.3 mm wide; silique 1.1–1.3 mm wide; embryo cotyledons with violet-purple veinsporphyroneura
46Anthers maroon or violet47
Anthers cream, yellow, pale yellow, sometimes flushed maroon49
47Lateral pinnae up to 8, petiolules 3.0–20.0 mm long; seed 1.2–1.8 mm longcoronata
Lateral pinnae up to 4, petiolules 0.1–1.5 mm long; seed 0.8–1.3 mm long48
48Leaves glossy; terminal leaflet orbicular-reniform to deltoid-reniform, base strongly cordate to occasionally ± truncate; cauline leaves 2.0–3.5 mm widebasicola
Leaves matt; terminal leaflet orbicular-pandurate, weakly pandurate to orbicular-rhomboid, base obtuse, truncate or weakly cordate; cauline leaves up to 1.5 mm widedactyloides
49Petals >4.5 mm50
Petals <4.5 mm54
50Silique width >1.5 mm51
Silique width <1.5 mm52
51Leaves coriaceous, lateral pinnae 2–6; terminal pinna 3.0–7.0 mm long, 2.8–8.6 mm wide, 1–3 marginal hydathodes; petals 4.2–7.0 mm long, 0.9–2.0 mm wide; siliques 1.5–2.0 mm wide; seeds 1.0–1.4 mm long, reticulate cells conspicuously thick-walledeminentia
Leaves membranous, lateral pinnae 1–2(–4); terminal pinna 4.7–36.0 mm long, 5.5–33.0 mm wide, up to 11 marginal hydathodes; petals 7.5–9.0 mm long, 2.5–5.0 mm wide; siliques 1.8–2.5 mm wide; seeds 1.8–2.2 mm long, reticulate cells not obviously thick-walledpolyodontes
52Sepals 1.2–1.6 mm wide; ovules 22–28; silique style 0.7–1.2 mm long; seeds greenchlorina
Sepals 1.0–1.3 mm wide; ovules 8–20; silique style 1.2–8.5 mm long; seeds henna53
53Siliques 1.0–1.2 mm wide; style 1.8–8.5 mm long; seeds 0.9–1.2 mm widedolichostyla
Siliques 1.1–1.4 mm wide; style 1.2–2.5 mm long; seeds 0.6–0.9 mm widecoronata
54Sepals 0.3–0.7 mm wide55
Sepals 0.7–1.4 mm wide57
55Stamens 4 (rarely 5–6)hirsuta
Stamens 656
56Annual herb; petals 1.8–2.3 mm long, 0.4–0.9 mm wide; silique style 0.3–0.5 mm long; seeds 0.7–0.9 mm long, 0.5–0.6 mm wideocculta
Perennial herb; petals 3.0–4.5 mm long, 0.7–1.4 mm wide; silique style 1.2–1.4 mm long; seeds 1.2–1.4 mm long, 0.6–0.9 mm wideforsteri
57Leaves membranous; inflorescence slightly flexuoseflexuosa
Leaves coriaceous or subcoriaceous; inflorescence straight58
58Leaves sparsely hairy, green; silique 9.0–20.0 mm long, style 1.0–1.4 mm longsubcarnosa
Leaves glabrous, grey to grey-green; silique 21.0–34.0 mm long, style 0.7–1.0 mm longthalassica
 Recognition

The following two tables provide habitat (Table 2) and character (Table 3) that may help distinguish between the species.

Synopsis of some major habitat types occupied and characters shared by species of Cardamine in New Zealand. o, obligate; f, facultative.

Habitat

Species

Alpine scree

C. alalata, C. eminentia, C. glara, C. parvula, C. serpentina, C. thalassica

Alpine bluffs, bouldery rubble

​​​​C. alticolaC. basicola, C. corymbosa, C. dimidia, C. intonsa, C. megalantha, C. sciaphila

Alpine wetland, seepages

C. exigua, C. reptans

Forest, shrubland

C. chlorina, C. corymbosa, C. dolichostyla, C. forsteri, C. intonsa, C. polyodontes, C. unguiculus

Limestone or marble

C. basicola (o), C. bilobata (f), C. bisetosa (o), C. caesiella (o), C. coronata (o), C. grandiscapa (f), C. integra (o), C. verna (o)

Low-elevation bluffs

C. dilatata, C. pachyphylla

River gravels

C. corymbosa, C. glara, C. verna​​​​​​ 

Shrubland, herbfield (main New Zealand islands)

C. corymbosa, C. dolichostyla, C. forsteri, C. polyodontes, C. unguiculus, C. unicaulis

Shrubland, herbfield (subantarctic islands)

C. corymbosa, C. depressa subsp. depressa, C. depressa subsp. stellata, C. latior, C. subcarnosa

Ultramafic outcrops

C. dactyloides (o), C. eminentia (f), C. porphyroneura (o), C. serpentina (o)

Weedy habitats occupied by indigenous species

C. corymbosa, C. forsteri, C. grandiscapa, C. heleniae, C. unicaulis

Weedy habitats occupied by naturalised species

C. flexuosa, C. hirsuta, C. occulta, C. pratensis

Wetland, seepages (often ephemeral)

C. cubita, C. exigua, C. lacustris, C. mutabilis, C. panatoheaC. reptans, C. sinuatifolia, C. unguiculus, C. unicaulis

 Distribution

Cardamine is a worldwide genus that occurs on all continents with the exception of Antarctica (Al-Shehbaz et al. 2006).  The New Zealand species occur on the main islands (North, South and Stewart Islands), numerous inshore islands (e.g., Hauraki Gulf islands), as well as many of the offshore islands such as Chatham Islands and the subantarctic Campbell Islands and Auckland Islands (Heenan 2017). They occur from the coast to the alpine zone, and occupy wetlands, forests, scree and rock bluff habitats.

 Biostatus
Indigenous (Non-endemic)
Number of species in New Zealand within Cardamine L.
CategoryNumber
Indigenous (Endemic)41
Indigenous (Non-endemic)1
Exotic: Fully Naturalised4
Total46
 Hybridisation

Cardamine verna × C. corymbosa. A single plant has features that indicate it may be a putative hybrid between C. verna (female) and C. corymbosa (male) (CHR 618326). This plant was collected from Isolation Creek, Marlborough, where it was growing among other seedlings of C. verna and with C. corymbosa growing nearby; this putative hybrid plant was cultivated at Lincoln. A hybrid origin is indicated by features of C. corymbosa, such as corymbose inflorescence; patent hairs on sepals and leaf margins; and rounded cauline leaflets. Features of C. verna include large and robust growth habit; leaves upright to spreading, brown-green, slightly irregular in shape, up to 120 mm long and coriaceous; and the inflorescences branched, upright, and up to 200 mm high. In addition, using Alexander’s Differential Stain, pollen stainability of the putative hybrid was moderate (54.7%), whereas plants of the putative parents from this site have high percentages of pollen stainability (C. verna 98.6%; C. corymbosa 99.6%).

Cardamine corymbosa × C. verna. Three plants have features that indicate they may be putative hybrids between C. corymbosa (female) and C. verna (male) (e.g., CHR 618324–618325). These plants were collected from Isolation Creek, Marlborough, where they were growing among other plants of C. corymbosa and with C. verna growing nearby; the putative hybrid plants were cultivated at Lincoln. These plants resemble C. corymbosa in having a compact growth habit; leaves low and spreading and with patent marginal hairs; corymbose inflorescences; and apetalous flowers. Features of C. verna include the robust growth habit; leaves up to 140 mm long and the leaflets irregularly shaped; and inflorescences up to 180 mm long. Using Alexander’s Differential Stain, pollen stainability of the three putative hybrids was low to moderate (27.4%, 29.4%, and 62.1%), whereas plants of the putative parents from this site have close to 100% pollen stainability (see above).

 Bibliography
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Heenan, P.B. 2020: Cardamine. In: Wilton, A.D. (ed.) Flora of New Zealand - Seed Plants. Fascicle 6. Manaaki Whenua Press, Lincoln.
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