- Cardamine alalata
- Cardamine alticola
- Cardamine basicola
- Cardamine bilobata
- Cardamine bisetosa
- Cardamine caesiella
- Cardamine chlorina
- Cardamine coronata
- Cardamine corymbosa
- Cardamine cubita
- Cardamine dactyloides
- Cardamine depressa
- Cardamine dilatata
- Cardamine dimidia
- Cardamine dolichostyla
- Cardamine eminentia
- Cardamine exigua
- Cardamine flexuosa
- Cardamine forsteri
- Cardamine glara
- Cardamine grandiscapa
- Cardamine heleniae
- Cardamine hirsuta
- Cardamine integra
- Cardamine intonsa
- Cardamine lacustris
- Cardamine latior
- Cardamine magnifica
- Cardamine megalantha
- Cardamine mutabilis
- Cardamine occulta
- Cardamine pachyphylla
- Cardamine panatohea
- Cardamine parvula
- Cardamine polyodontes
- Cardamine porphyroneura
- Cardamine pratensis
- Cardamine reptans
- Cardamine sciaphila
- Cardamine serpentina
- Cardamine sinuatifolia
- Cardamine subcarnosa
- Cardamine thalassica
- Cardamine unguiculus
- Cardamine unicaulis
- Cardamine verna
- = Iti Garn.-Jones & P.N.Johnson, New Zealand J. Bot. 25: 603 (1987)
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.
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.
1 | Fruit a silicle or silique, <4.0 mm long | 2 |
Fruit a silique, >6.0 mm long | 3 | |
2 | Leaves broadly orbicular, <2.2 mm long; 2 stamens, bent at distal end; silique <4.0 mm long | cubita |
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 long | lacustris | |
3 | Inflorescence decumbent or prostrate, axillary rosettes present in nodes | panatohea |
Inflorescence erect or spreading, axillary rosettes absent from nodes | 4 | |
4 | Leaves 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 coriaceous | 5 |
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 membranous | 13 | |
5 | Anthers maroon or yellow and flushed maroon | 6 |
Anthers yellow | 8 | |
6 | Sepals 0.8–1.1 mm wide; petals 4.0–7.0 mm long; silique style 0.3–0.5 mm long | dimidia |
Sepals 1.3–1.8 mm wide; petals 7.0–11.0 mm long; silique style 1.0–3.5 mm long | 7 | |
7 | Leaves 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 long | bilobata |
Leaves grey-green to green; filaments conspicuously dilated at base, 0.35–0.65 mm wide; silique style 1.0–1.2 mm long | dilatata | |
8 | Leaves always simple | 9 |
Leaves simple and pinnatisect or pinnatifid | 10 | |
9 | Leaves 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 long | alticola |
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 long | sciaphila | |
10 | Leaves 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 ascending | mutabilis |
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 erect | 11 | |
11 | Leaves 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 Islands | depressa |
Leaves coriaceous, lamina surface glabrous; inflorescence usually elongated and held well above the leaves, flowers distributed along inflorescence, plants of South Island | 12 | |
12 | Leaves 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 long | pachyphylla |
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 long | dimidia | |
13 | Leaves with axillary hydathodes | 14 |
Leaves without axillary hydathodes | 20 | |
14 | Siliques glabrous | 15 |
Siliques hairy | 18 | |
15 | Leaves coriaceous, terminal leaflet 3.0–7.7 mm long and 2.8–8.6 mm wide; seed wing present, reticulate cells conspicuously thick-walled | eminentia |
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-walled | 16 | |
16 | Lateral 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 wide | megalantha |
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 wide | 17 | |
17 | Leaf 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 wide | coronata |
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 wide | forsteri | |
18 | Terminal 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 long | bisetosa |
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 long | 19 | |
19 | Leaf 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 wide | coronata |
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 wide | forsteri | |
20 | Seeds winged | 21 |
Seeds not winged | 23 | |
21 | Leaves membranous; silique 1.5–2.3 mm wide; seed oblong to oblong-elliptic, green to green-brown, wing at apex | unguiculus |
Leaves coriaceous; silique 1.1–1.9 mm wide; seed orbicular, broadly oblong to oblong, henna, wing at apex and lateral margin | 22 | |
22 | Leaves 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 long | alalata |
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 long | verna | |
23 | Siliques hairy | 24 |
Siliques glabrous | 28 | |
24 | Silique hairs sparse, patent, 0.35–0.6 mm long | hirsuta |
Silique hairs moderate to dense, appressed to spreading, up to 0.35 mm long | 25 | |
25 | Petals 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 long | coronata |
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 long | 26 | |
26 | Leaves 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 maroon | bisetosa |
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 yellow | 27 | |
27 | Sepals 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 long | intonsa |
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 long | forsteri | |
28 | Inflorescence corymbose | 29 |
Inflorescence racemose | 40 | |
29 | Petals >6.5 mm long | reptans |
Petals <6.5 mm long | 30 | |
30 | Petals <3.5 mm long | 31 |
Petals 3.5–6.5 mm long | 33 | |
31 | Leaves 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 wide | heleniae |
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 wide | 32 | |
32 | Leaves 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 long | exigua |
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 long | parvula | |
33 | Terminal leaflet longer than wide, base attenuate to cuneate, occasionally obtuse | 34 |
Terminal leaflet similar length to width, base obtuse, truncate or cordate | 35 | |
34 | Leaves 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 long | integra |
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 long | mutabilis | |
35 | Leaves coriaceus | 36 |
Leaves membranous | 37 | |
36 | Leaves 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 long | serpentina |
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 long | glara | |
37 | Sepals 1.1–1.6 mm long; silique 8.0–13.5 mm long | unicaulis |
Sepals 1.3–3.0 mm long; silique 12.0–32.0 mm long | 38 | |
38 | Inflorescence 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 wide | grandiscapa |
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 wide | 39 | |
39 | Leaves 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 long | sinuatifolia |
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 long | corymbosa | |
40 | Lateral pinnae ≥10 | 41 |
Lateral pinnae ≤8 | 43 | |
41 | Terminal 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 wide | latior |
Terminal pinna 2–25 mm long, 2–27 mm wide; inflorescence up to 350 mm long; silique 1.0–1.7 mm wide | 42 | |
42 | Leaf 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 long | pratensis |
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 long | flexuosa | |
43 | Lateral pinnae ≤2 | 44 |
Lateral pinnae ≥3 | 46 | |
44 | Leaves coriaceous, grey-green to blue-grey, broadly ovate or broadly orbicular-deltoid, base truncate; sepals sparsely hairy | caesiella |
Leaves membranous, green, pandurate, pandurate-orbicular or rhomboid-pandurate, base cordate; sepals glabrous | 45 | |
45 | Sepals 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 veins | megalantha |
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 veins | porphyroneura | |
46 | Anthers maroon or violet | 47 |
Anthers cream, yellow, pale yellow, sometimes flushed maroon | 49 | |
47 | Lateral pinnae up to 8, petiolules 3.0–20.0 mm long; seed 1.2–1.8 mm long | coronata |
Lateral pinnae up to 4, petiolules 0.1–1.5 mm long; seed 0.8–1.3 mm long | 48 | |
48 | Leaves glossy; terminal leaflet orbicular-reniform to deltoid-reniform, base strongly cordate to occasionally ± truncate; cauline leaves 2.0–3.5 mm wide | basicola |
Leaves matt; terminal leaflet orbicular-pandurate, weakly pandurate to orbicular-rhomboid, base obtuse, truncate or weakly cordate; cauline leaves up to 1.5 mm wide | dactyloides | |
49 | Petals >4.5 mm | 50 |
Petals <4.5 mm | 54 | |
50 | Silique width >1.5 mm | 51 |
Silique width <1.5 mm | 52 | |
51 | Leaves 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-walled | eminentia |
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-walled | polyodontes | |
52 | Sepals 1.2–1.6 mm wide; ovules 22–28; silique style 0.7–1.2 mm long; seeds green | chlorina |
Sepals 1.0–1.3 mm wide; ovules 8–20; silique style 1.2–8.5 mm long; seeds henna | 53 | |
53 | Siliques 1.0–1.2 mm wide; style 1.8–8.5 mm long; seeds 0.9–1.2 mm wide | dolichostyla |
Siliques 1.1–1.4 mm wide; style 1.2–2.5 mm long; seeds 0.6–0.9 mm wide | coronata | |
54 | Sepals 0.3–0.7 mm wide | 55 |
Sepals 0.7–1.4 mm wide | 57 | |
55 | Stamens 4 (rarely 5–6) | hirsuta |
Stamens 6 | 56 | |
56 | Annual 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 wide | occulta |
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 wide | forsteri | |
57 | Leaves membranous; inflorescence slightly flexuose | flexuosa |
Leaves coriaceous or subcoriaceous; inflorescence straight | 58 | |
58 | Leaves sparsely hairy, green; silique 9.0–20.0 mm long, style 1.0–1.4 mm long | subcarnosa |
Leaves glabrous, grey to grey-green; silique 21.0–34.0 mm long, style 0.7–1.0 mm long | thalassica |
The following two tables provide habitat (Table 2) and character (Table 3) that may help distinguish between the species.
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.
Category | Number |
---|---|
Indigenous (Endemic) | 41 |
Indigenous (Non-endemic) | 1 |
Exotic: Fully Naturalised | 4 |
Total | 46 |
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).