Cardamine pratensis, Linnaeus, 1753

Heenan, Peter B., 2017, A taxonomic revision of Cardamine L. (Brassicaceae) in New Zealand, Phytotaxa 330 (1), pp. 448-450 : 448-450

publication ID	https://doi.org/ 10.11646/phytotaxa.330.1.1
persistent identifier	https://treatment.plazi.org/id/03B7878D-3010-FF13-FF1F-D77505919623
treatment provided by	Felipe
scientific name	Cardamine pratensis
status

Treatment

Cardamine pratensis View in CoL Toothed-leaved form ( Fig. 74B View FIGURE 74 )

Leaves with 16–24 lateral pinnae, broadly elliptic to elliptic-ovate and margin toothed, base cuneate to obtuse. Cauline leaves narrower, smaller and with fewer pinnae than rosette leaves, entire or with 1–4 pairs of teeth; terminal pinnae up to 17.0 × 1.8 mm; lateral pinnae up to 15.0 × 1.2 mm.

HEENAN

Distribution and habitats:—Naturalised in New Zealand. Known from Hauraki Plains, North Auckland, where it occurs in swamps and grassland.

Representative specimens:— NEW ZEALAND. North Auckland. Te Awaiti Canal, Hauraki plains, 16 October 1945, A. J. Healy s.n., CHR 45939; Matataki, Waihou River , Hauraki plains , 17 October 1945, A. J. Healy s.n., CHR 45938; Waihou River , Mangati, 3 December 1959, R. Mason 7444, CHR 113261; Waihou River , Pukahu Road, R. Mason 7336, 1 December 1959, CHR 113076; Komata Road, Hauraki plains , 1 December 1959, R. Mason 7331, CHR 113071 .

Notes:—In New Zealand the tooth-leaved form of C. pratensis is only known from the Hauraki Plains, North Auckland. It is distinguished from the more typical sinuate margin/entire-leaved form of C. pratensis by more numerous leaflets that are prominently toothed, the silique is slightly more slender and with a longer style, and the seeds are oblong rather than being rounded.

Cardamine reptans Heenan , spec. nov. ( Fig. 76 View FIGURE 76 )

Distinguished from C. megalantha by its membranous leaves with indistinct marginal hydathodes, smaller terminal leaflet and in having several lateral leaflets, leaflets often with indistinct petiolules and lacking axillary hydathodes, and short inflorescences.

Holotype:— NEW ZEALAND. Otago Land District, Dunstan Mts., Fairfax Spur , near leaning Rock , 9 December 1997, P. B. Heenan s.n. (CHR 514169!).

HEENAN NEW ZEALAND CARDAMINE REVISION

Phytotaxa 330 (1) © 2017 Magnolia Press • 129 Perennial herb, single rosette or with lateral, prostrate or decumbent, often with rhizomatous stems. Leaves up to 95 mm long, pinnatisect; lamina 5.0–35.0 × 3.5–34.0 mm, green, coriaceous, glabrous on abaxial surface, glabrous or sparsely to moderately hairy on adaxial surface and petiole; hairs up to 0.4mm long, spreading to patent; petiole up to 75 mm long. Terminal pinna 2.5–13.0 × 2.0– 14.8 mm, simple, orbicular-rhomboid, rhomboid, broadly

HEENAN elliptic-rhomboid to broadly elliptic; margin entire to shallowly sinuate, with 2–4 indistinct hydathodes; apex obtuse with a ± distinct hydathode; base attenuate to obtuse, occasionally truncate. Lateral pinnae 2–6, 1.0–13.0 × 0.6–8.3 mm, orbicular to broadly elliptic-orbicular; apex obtuse; base attenuate; petiolule sessile or up to 5.0 mm long. Cauline leaves absent. Inflorescence corymbose, each corymb 2–3-flowered or flowers solitary; peduncle up to 45 mm long, 0.3–0.4 mm diam. at base, spreading to ascending, glabrous. Pedicels 6.0–145.0 mm long, 0.3–0.4 mm diam., glabrous. Sepals 2.2–3.4 × 1.0– 1.8 mm, elliptic-oblong to broadly elliptic-obovate, saccate, green or reddish-brown, glabrous or occasionally sparsely hairy, margin white and membranous, apex obtuse, base truncate. Petals 6.5–9.7 × 3.0– 5.6 mm, white, limb obovate; apex obtuse; base attenuate to cuneate, tapering to a 2.0– 2.5 mm long claw. Stamens 6; median filaments 4, 3.6–5.2 mm long; lateral filaments 2, 2.0– 3.7 mm long; anthers 0.9–1.0(–2.2) mm long, cream to pale yellow, when dehiscent held at a similar height to or slightly below the stigma. Ovary 5.0– 6.4 mm long, 0.5–0.9 mm diam., ± terete, green, glabrous; ovules 28–36; style (1.2–)2.5–3.0 mm long, ± terete; stigma 0.35–0.45 mm diam. Siliques 11.0–15.0 × 1.5–1.7 mm, glabrous, style 1.6–2.0 mm long; valves green at maturity and when dehiscent; replum 0.2–0.3 mm wide. Seeds 1.3–1.7 mm long, 0.8–1.1 mm wide, 0.2–0.4 mm thick, oblong to broadly oblong, henna to henna-green; wing absent. FL Dec–Feb; FT Jan–Feb.

Representative specimens:— NEW ZEALAND. Otago. Kurow Mts , January 1931, A. Wall s.n., CHR 329522 ; St Bathans Range , 10 January 1972, C. D. Meurk s.n., OTA 34556 About OTA ; St Mary’s Range , 5 February 1998, P. B. Heenan s.n., CHR 546244 ; St Mary’s Range , 19 December 1987, B. D. Rance s.n., CHR 459551 ; Dunstan Mountains , 13 December 2011, J. Barkla s.n., CHR 619272 ; Mt Pisa, 1885-1890, D. Petrie s.n., AK 4438; Pisa Range , 25 February 1997, P. B. Heenan s.n. & M. I. Dawson, CHR 511718 ; Old Man Range , 30 December 1963, A. F. Mark s.n., OTA 9092 About OTA ; Old Man Range , 30 December 1963, A. F. Mark s.n., CHR 9092 / CHR 9093; Old Man Range , 27 December 1968, A. F. Mark & N. M. Adams s.n., OTA 7014 About OTA ; Gorge Creek, Old Man Range , 12 February 1966, J. Wells s.n., OTA 61147 About OTA ; Old Man Range , 24 February 1997, P. B. Heenan s.n. & M. I. Dawson, CHR 511701 / CHR 511684; Old Man Range , 30 December 2016, P. B. Heenan s.n., CHR 643940 ; Remarkables , December 1991, A. P. Druce 1477, CHR 472069 ; Mt Cardrona, Crown Range , February 1992, A. P. Druce 1476, CHR 472068 ; Mt Cardrona, Crown Range , January 1992, A. P. Druce 1478, CHR 472070 ; Mt Earnslaw , 1 March 1997, D. Glenny 6881, CHR 530557 . Southland. Lake Laura, Garvie Mountains , 17 February 2004, B. D. Rance s.n., CHR 595711 . Cultivated. ex Old Man Range, Otago, cultivated experimental nursery, Landcare Research , 17 September 2010, P. B. Heenan s.n., CHR 618390 /CHR 618391.

Distribution and habitats:— New Zealand, endemic. Cardamine reptans is known from the schist mountains of Otago, including St Mary’s Range, St Bathans Range, Dunstan Mountains, Old Man Range, Pisa Range, Remarkables and Crown Range, and Southland (Garvie Mountains) ( Fig. 68 View FIGURE 68 ). It occurs in variety subalpine and alpine habitats such as seepages, damp stream margins, moist hollows, bog margins, and wet hollows in alpine cushion vegetation.

Conservation status:— Cardamine reptans is assessed as having a conservation status of At Risk—Naturally Uncommon, with the qualifier Data Poor ( Townsend et al. 2008). The qualifier Data Poor is applied as additional information on the number and size of the populations is required.

Etymology:—The specific epithet reptans (Latin: creeping) refers to the growth habit.

Notes:—Previously known by the tagname C. “corymbosa large flower”.

Cardamine sciaphila Heenan , spec. nov. ( Fig. 77 View FIGURE 77 )

Distinguished from C. alticola by its glossy, narrow and coriaceous leaves that are often hairy, shorter petioles, and its petals being 2.1–3.2 mm long.

Holotype:— NEW ZEALAND. Otago Land District, Dunstan Mts , among grasses around rock tors, 9 December 1997, P. B. Heenan s.n. (CHR 514168!).

Perennial herb, single rosette or with short lateral branches, stem and branches c. 2 mm diam. Leaves up to 17(–45) mm long, simple, glabrous or hairy; lamina 5–20 × 2–5 mm, narrowly obovate to spathulate, glossy, coriaceous, green, brown-green to bronze, glabrous on abaxial surface, glabrous or sparsely to moderately hairy on margin and adaxial surface; apex obtuse, often with a distinct hydathode; base cuneate to attenuate, grading into petiole; hairs unicellular, straight or curved, transparent. Petiole up to 12(–25) mm long, plano-convex, usually glabrous although sometimes 1–3 hairs on margin. Cauline leaves occasional, similar to rosette leaves but smaller. Inflorescence corymbose with 2–8 flowers, flowers sometimes solitary, peduncle up to 30(–70) mm long, 0.7–0.8 mm

HEENAN NEW ZEALAND CARDAMINE REVISION

Phytotaxa 330 (1) © 2017 Magnolia Press • 131 diam. at base, upright, glossy, glabrous, green to brown-green. Pedicels up to 30 mm long, 0.5 mm diam., flexuose. Sepals 1.6–1.8 × 0.7–1.0 mm, oblong to oval, ± saccate, green, glabrous or with occasional simple hairs, margin white and membranous, apex obtuse, base truncate. Petals 2.1–3.2 × 1.2–1.3 mm, white, limb obovate to oblong; apex obtuse; base attenuate, tapering into the ± indistinct claw. Stamens 4–6; filaments 1.2–1.3 mm long; anthers

HEENAN

0.4–0.5 mm long, yellow, held at a similar height to or slightly overtopping the stigma. Ovary 1.8–2.0 mm long, c. 0.6 mm diam., ± terete, green, glabrous; ovules 13–14 in each locule; style 0.3–0.4 mm long, ± terete; stigma 0.4–0.5 mm diam. Siliques 7–14 × 1.4–1.5 mm, erect, glabrous, valves green at maturity, style 0.7–0.8 mm long. Seeds 1.0– 1.1 mm long, 0.8–0.9 mm wide, 0.3–0.5 mm thick, oblong, orbicular-oblong to orbicular, pale yellow; wing absent. FL Dec–Jan; FT Jan–Mar.

Representative specimens:— NEW ZEALAND. Otago. Dunstan Range, 24 February 1997, P. B. Heenan s.n. & M. I. Dawson, CHR 511706/CHR 511691; Mt Pisa , Pisa Range, March 1989, A. P. Druce s.n., CHR 395191; Pisa Range, 24 January 2006, J. Barkla s.n., CHR 580889 . Cultivated. ex Dunstan Mts, Otago, cultivated experimental nursery, Landcare Research , 2 March 1998, P. B. Heenan s.n. & A. Mitchell, CHR 515296; ex Dunstan Mts , Otago, cultivated experimental nursery, Landcare Research, 5 June 1998, P. B. Heenan s.n., CHR 515526 .

Distribution and habitats:— New Zealand, endemic. Cardamine sciaphila is known from the highest parts of the Dunstan Mountains (1600 m) and Pisa Range (1620–1850 m) in Central Otago ( Fig. 78 View FIGURE 78 ). It occurs among lowgrowing herbs and grasses on ledges and crevices of rock tors, outcrops, and bluffs, showing a clear preference for shaded sites. Plants are often difficult to find unless in flower as they are often buried among other herbaceous vegetation.

Etymology:—The specific epithet sciaphila (Latin: shade-loving) refers to the shaded habitat that is preferred by the species.

Conservation status:— Cardamine sciaphila is assessed as having a conservation status of Threatened, Nationally Critical (A1), with the qualifier Data Poor ( Townsend et al. 2008). The qualifier Data Poor is applied

HEENAN NEW ZEALAND CARDAMINE REVISION

Phytotaxa 330 (1) © 2017 Magnolia Press • 133 since there is suitable alpine habitat for this species on the plateau-mountains of Central Otago and field survey is likely to locate additional populations. This conservation assessment is consistent with that of de Lange et al. (2013, as C. “Pisa”).

Notes:—Plants from the Dunstan Mountains have a conspicuously hairy leaf lamina whereas the lamina of plants from the Pisa Range is very sparsely hairy, glabrate or glabrous. Few flowers were available for study, but a plant from the Pisa Range (CHR 580889) had two flowers that had only four stamens whereas plants from Dunstan Mountains had six stamens. Further taxonomic research on variation of the leaf and flower characters is required.

Previously known by the tagname C. “Pisa Range” (e.g., de Lange et al. 2013, p. 65).

Cardamine serpentina Heenan , spec. nov. ( Fig. 79 View FIGURE 79 )

Distinguished from C. glara by its smaller and compact growth habit, smaller green-brown to purple-brown leaves, smaller terminal leaflet with a more distinct cordate base, fewer lateral pinnae, smaller sepals, and shorter siliques.

Holotype:— NEW ZEALAND. Southland Land District, West Dome , ultramafic scree, 27 January 1999, P. B. Heenan s.n. (CHR 546249!).

Perennial rosette herb, stem and branches c. 2 mm diam. Leaves up to 40 mm long, simple or pinnatisect; lamina 3–13 × 3–8 mm, green-brown to purple-brown, coriaceous, glabrous; terminal pinna 2.0–6.0 × 1.8–8.0 mm, orbicular-reniform to deltoid-reniform, apex obtuse with a small but distinct hydathode; base truncate to cordate. Lateral pinnae 1–2 or absent, 0.8–3.2 × 0.8–2.3 mm, orbicular, petiolule up to 1.2 mm long; petiole up to 32 mm long, plano-convex, glabrous. Cauline leaves sometimes subtending lower pedicels, narrowly obovate, up to 13 mm long. Inflorescence corymbose to corymbose-racemoase, with 1–3 lateral corymbs, each corymb 2–8- flowered, flowers sometimes solitary, peduncle up to 100 mm long, 0.6–0.9 mm diam. at base, upright, glabrous, green-brown to purple-brown. Pedicels up to 36 mm long, 0.3–0.5 mm diam., flexuose, ascending to divaricate. Sepals 1.8–2.4 × 1.0– 1.8 mm, broadly oblong to ovate, weakly saccate, green, often flushed red-brown, glabrous, margin white and membranous, apex obtuse, base truncate. Petals 4.5–5.0 × 2.6–3.0 mm, white, limb obovate; apex obtuse; base attenuate, tapering into the ± indistinct claw. Stamens 6; median filaments 4, 2.7–3.1 mm long, lateral filaments 2, 2.9–3.5 mm long; anthers 0.6–0.7 mm long, yellow, overtopping the stigma. Ovary 2.7–3.2 mm long, 0.5–0.6 mm diam., ± terete, green, glabrous; ovules 13–14 in each locule; style 0.3–0.5 mm long, ± terete; stigma 0.3–0.4 mm diam. Siliques 12–20 × 0.9–1.2 mm, erect, glabrous, valves green-brown to purple-brown at maturity, style 0.8–1.8 mm long. Seeds 1.0– 1.2 mm long, 0.8–0.9 mm wide, 0.3–0.4 mm thick, orbicular to orbicular-oblong, orange-brown; wing absent. FL Dec–Feb; FT Dec–Apr.

Representative specimens:— NEW ZEALAND. Southland. West Dome A. P. Druce s.n., CHR 624630 ; West Dome, 9 January 1995, P. J. de Lange s.n., AK 231673; West Dome , 28 January 1998, P. B. Heenan s.n., CHR 514964 ; West Dome , 27 December 1999, P. B. Heenan s.n., CHR 536721 ; West Dome , 15 December 2009, B. Rance & G. Rogers s.n., CHR 616877 ; Mt Cerberus, Livingstone Mts , 4 January 1984, W. G. Lee 48, CHR 405205 ; Mt Cerberus, Livingstone Mts , 27 January 1998, P. B. Heenan s.n., CHR 514971 ; Mt Cerberus, Livingston Mts , 5 February 2007, M. Thorsen 51/07, CHR 591782 ; Black Ridge Quarry , 14 April 1975, W. G. Lee s.n., OTA 35930 About OTA ; Black Ridge Quarry , 1 December 1978, P. N. Johnson s.n., CHR 320241 ; Black Ridge Quarry , 15 April 2004, P. B. Heenan s.n., CHR 573472 . Cultivated. ex Bald Hill, Southland, cultivated experimental nursery, Landcare Research , 17 September 2010, P. B. Heenan s.n., CHR 618392 .

Distribution and habitats:— New Zealand, endemic. Cardamine serpentina is restricted to Southland where it occurs at West Dome, Black Ridge Quarry, Bald Hill, and Livingstone Mountains ( Fig. 78 View FIGURE 78 ). It grows on scree and loose stones/rubble around outcrops derived from ultramafic rocks.

Conservation status:— Cardamine serpentina is assessed as having a conservation status of Threatened, Nationally Vulnerable (B1) ( Townsend et al. 2008). This assessment differs from the At Risk—Naturally Uncommon conservation listing by de Lange et al. (2013, as C. “West Dome”). This revised assessment of C. serpentina is consistent with its small population sizes, geographic restriction and specific habitat (see Townsend et al. 2008, p. 24).

Etymology:—The specific epithet serpentina (Latin: serpentine rock) refers to the base rock and substrate the species occurs on.

Notes:—Previously known by the tag name C. “West Dome” (e.g., de Lange et al. 2013, p. 68).

HEENAN HEENAN NEW ZEALAND CARDAMINE REVISION

Cardamine sinuatifolia Heenan , spec. nov. ( Figs. 8 View FIGURE 8 , 80 View FIGURE 80 )

Distinguished from C. corymbosa by its light green leaves with a sinuate margin, larger sepals and petals, and longer silique style.

Holotype:— NEW ZEALAND. Canterbury, Torlesse Range, Foggy Peak , damp banks of stream, 9 December 2009, P. B. Heenan s.n. (CHR 616910!).

Perennial herb, single rosette or with short lateral branches. Leaves up to 90 mm long, simple or pinnatisect, upright to spreading; lamina 4.0–30.0 × 2.5–15.0 mm, light green, membranous, usually glabrous on abaxial and adaxial surfaces, glabrous or sparsely hairy on margin and petiole; petiole up to 70 mm long, green and usually purple-red in lower half, glabrous to sparsely hairy, hairs 0.3–0.5 mm long, spreading to patent. Terminal pinna 5.0–18.0 × 5.0–15.0 mm, simple, oblong-orbicular, broadly elliptic-orbicular, elliptic-oblong to deltoid-oblong; margin usually shallowly sinuate to occasionally entire, hydathodes indistinct; apex obtuse, hydathode indistinct; base truncate, obtuse, to weakly cordate. Lateral pinnae 1–2(–4), 2.0–8.0 × 1.5–6.0 mm, orbicular, broadly elliptic-orbicular, deltoid-orbicular to rhomboid-orbicular; margin entire, occasionally weakly lobed, hydathodes inconspicuous; apex obtuse; base truncate to cuneate, often oblique; petiolule 0.2–1.0 mm long, often appearing ±sessile. Cauline leaves similar to rosette leaves but smaller, terminal pinna often pinnatifid, pinnae bases attenuate to truncate; terminal pinna up to 10.0 × 6.0 mm, lateral pinnae up to 8.0 × 4.5 mm. Inflorescence with solitary flowers or 1–3 corymbs, corymbs 2–6-flowered; peduncle up to 100 mm long, 0.5–1.0 mm diam. at base, spreading to ascending, glabrous. Pedicels 15.0–115.0 mm long, 0.3–0.7 mm diam., terete, glabrous. Sepals 1.8–2.2 × 0.7–1.2 mm, elliptic-oblong, saccate, green or red-brown; glabrous to sparsely hairy; margin white and membranous, apex obtuse, base truncate. Petals 1–4, or absent; 4.5–5.5 × 2.1–3.0 mm, white, limb obovate; apex obtuse; base cuneate, tapering to a 1.0– 1.5 mm long claw. Stamens 4–6; median filaments 2–4, 2.0– 2.6 mm long; lateral filaments 2, 1.6–2.2 mm long; anthers 0.5–0.6 mm long, cream to pale yellow, when dehiscent held at a similar height to or slightly below the stigma. Ovary 2.6–2.9 mm long, 0.4–0.6 mm diam., ± terete, green, glabrous; ovules 20–20; style c. 0.2 mm long, ± terete; stigma 0.4–0.5 mm diam., white. Siliques 18.0–25.0 × 1.0– 1.2 mm, glabrous, style 0.2–0.4 mm long; valves green at maturity and when dehiscent; replum 0.4–0.5 mm wide. Seeds 0.9–1.2 mm long, 0.5–0.7 mm wide, 0.3–0.4 mm thick, oblong to broadly oblong, henna; wing absent. FL Jan–Dec; FT Jan–Mar.

Representative specimens:— NEW ZEALAND. Nelson. Lake Henderson, Peel Range , March 1982, A. P. Druce s.n., CHR 369975 . Canterbury. Foggy Peak, 9 December 2009, P. B. Heenan s.n., CHR 617181/CHR 616911; Broken River Ski Club, 25 January 1964, herbarium J. Thompson s.n., CHR 522525 . Otago. Twelve Mile Creek, 26 February 1997, P. B. Heenan s.n. & M. I. Dawson, CHR 511934; Lammerlaw Range , 16 December 2009, J. Barkla s.n., CHR 610176 . Cultivated. ex Lake Henderson, Nelson, cultivated experimental nursery, Landcare Research , 3 December 2009, P. B. Heenan s.n., CHR 618384; ex Foggy Peak, Canterbury, cultivated experimental nursery, Landcare Research, 3 December 2009, P. B. Heenan s.n., CHR 618376; ex Broken River skifield, Craigieburn Range, Canterbury, cultivated experimental nursery, Landcare Research, 6 March 2014, P. B. Heenan s.n., CHR 636124 .

Distribution and habitats:— New Zealand, endemic. Cardamine sinuatifolia is known from Nelson, Canterbury and Otago in the South Island ( Fig. 78 View FIGURE 78 ). Although currently known from few collections, the wide geographic range of C. sinuatifolia in the South Island suggests it is likely to be more common. Furthermore, its preferred habitat of damp sites on lake margins and stream banks is reasonably common.

Conservation status:— Cardamine sinuatifolia is assessed as having a conservation status of At Risk—Naturally Uncommon, with the qualifiers Data Poor and Sparse ( Townsend et al. 2008). The qualifier Data Poor is applied as there is a lack of information on the number and size of the populations. Sparse is applied to reflect the current distribution, but further populations will almost certainly be located since its habitats are common throughout its distributional range.

Etymology:—The specific epithet sinuatifolia (Latin: wavy leaves) refers to the leaves of this species usually being wavy on the margin.

Notes:—Distinguished by usually simple leaves, although occasionally they have 1–2 lateral pinnae, light green colour, margin with inconspicuous hydathodes, and purple-red petiole bases and stems. A notable feature of the terminal leaflets is their irregular shape, a tendency to be elliptic or oblong, and the weakly sinuate margin.

HEENAN HEENAN NEW ZEALAND CARDAMINE REVISION

Phytotaxa 330 (1) © 2017 Magnolia Press • 137 Cardamine subcarnosa (Hook.f.) Allan (1961: 184) ≡ Cardamine hirsuta var. subcarnosa ( Hooker 1844: 5) ≡ Cardamine glacialis var. subcarnosa (Hook.f.) Schulz (1903: 542) . Lectotype (fide Heenan 2008):— NEW ZEALAND. Flora of Campbell Islands, Antarctic Expedition, 1839–1843, J. D. Hooker (BM!). ( Fig. 81 View FIGURE 81 )

Perennial herb, caespitose, forming an open rosette. Leaves pinnatisect, up to 100(–140) mm long, green, glossy, subcoriaceous, lamina and petiole margins sparsely ciliate, glabrate, or occasionally glabrous; petiole 20–40(–60) mm long, 1.5–6.0 mm wide, winged and sheathing at base; hydathodes inconspicuous to prominent. Terminal pinna 5–25 × 5–20 mm, orbicular to broadly elliptic, with inconspicuous lateral lobes or shallowly toothed, apex obtuse to rounded, base cuneate, obtuse or ± truncate. Lateral pinnae 5–7, 3–22 × 2.5–12 mm, usually in pairs although proximal leaflets occ. alternate, usually not overlapping except occasionally the uppermost pair and the terminal, broadly elliptic, elliptic to obovate, shallowly toothed or entire; petiolules up to 10 mm long, although sometimes ± absent, apex obtuse to rounded, base cuneate, obtuse or ± truncate. Cauline leaves subtending pedicels, although sometimes absent on upper pedicels; lower leaves similar to rosette leaves, but with fewer and narrower leaflets, becoming smaller in all parts; upper leaves 1.7–6.5 × 0.3–0.9 mm, increasingly linear, simple. Inflorescence 50–150(–300) mm long, 1.2–1.6 mm diam. at base, glabrous, usually elongating after flowering, upright to ascending, racemose, flowers distant in upper half. Pedicels 2.0– 12 mm long, 0.5–0.8 mm diam, erecto-patent to spreading. Sepals 1.3–2.4 × 0.7–1.2 mm, oblong to elliptic, glabrous, green or purple, margin white and membranous, apex obtuse to rounded, base truncate. Petals 2.2–4.5 × 0.6–1.3 mm, white, pink or purple, usually purple veined, limb obovate; apex obtuse to rounded; base cuneate to attenuate, tapering to a ± indistinct claw, claw up to 0.5 mm long. Stamens 6; median filaments 4, 1.9–2.2 mm long; lateral filaments 2, 1.6–1.9 mm long; anthers 0.3–0.4 mm long. Ovary 3.2–3.5 mm long, 0.5–0.7 mm diam., terete, glabrous; stigma 0.3–0.5 mm diam. Siliques 9–20 × 0.9–1.3 mm, not crowded, erecto-patent to spreading, style 1.0– 1.4 mm long; replum 0.3–0.4 mm wide. Seeds 0.8–1.4 mm long, 0.5–0.9 mm wide, 0.3–0.5 mm thick, orbicular-oblong or oblong, red-brown. FL Nov–Feb; FT Dec–Feb.

Representative specimens:— NEW ZEALAND. Campbell Island: unknown locality, November 1907, R. M. Laing s.n., CHR 329524/CHR 329525; Lyall Ridge, 4 December 1944, R. L. Oliver s.n., CHR 277037; Mt Azimuth, 9 February 1946, J. H. Sorensen s.n., CHR 277039; Azimuth, February 1945, J. F. Findlay s.n., CHR 49784/CHR 49785; Mt St Col, 23 December 1946, W. B. B [rockie] s.n., CHR 233470; Faye Ridge, February 1985, G. A. Taylor s.n., AK 295669; Campbell Island, November 1907, M. Laing s.n., AK 4442; Mt Honey, 23 January 1961, E. J. Godley s.n., CHR 118007; St. Col, January 1961, F. Fisher s.n., CHR 156443; Lyall Ridge, 6 January 1961, V. D. Zotov s.n., CHR 117891; Lyall Ridge, 6 January 1961, F. J. Fisher s.n., CHR 117897; Mt Azimuth, 18 January 1961, E. J. Godley s.n., CHR 118000; N. E. Valley, 1975, P. Wilson s.n., CHR 311529; St Col Peak, 18 December 1975, D. R. Given 8981, CHR 303741; Moubray Hill, 22 December 1975, D. R. Given 9016, CHR 284791; Yvon Villarceau Peak, 30 January 1976, D. R. Given 9318, CHR 323126; Mount Azimuth, 7 January 1976, D. R. Given 9116, CHR 303801; Mount Azimuth, D. R. Given 9117, 7 January 1976, CHR 303800; Mount Azimuth, 7 January 1976, D. R. Given 9166, CHR 303823; Mount Honey, 15 January 1976, D. R. Given 9210, CHR 303841; Mount Honey, 15 January 1976, D. R. Given 9209, CHR 303842; Mt Honey, 9 January 2011, S. J. Wagstaff 112, CHR 624024; Mount Dumas, 10 January 1976, D. R. Given 9132, CHR 303783; unknown locality, 1943, J. H. Sorenson s.n., CHR 277036; N. W. Bay, 3 February 1947, W. B. B[rockie] s.n., WELT 27973; unknown locality, November-December 1975, B. May s.n., AK 162785.

Distribution and habitats:— New Zealand, endemic. Cardamine subcarnosa is known only from Campbell Island. It grows in damp places on scree, peat, grassland, and rock crevices. Noted by Meurk et al. (1994) as being most common in herbfields, tall Marsippospermum rushlands, and fellfields of the summit tundra zone.

Conservation status:— Cardamine subcarnosa is assessed as having a conservation status of At Risk—Naturally Uncommon, with the qualifiers Data Poor, Island Endemic and One Location ( Townsend et al. 2008). The qualifier Data Poor is applied as additional information is required on the number and size of the populations. This conservation assessment is consistent with that of de Lange et al. (2013).

Etymology:—The specific epithet subcarnosa (Latin: sub a little; carnosa fleshy) probably refers to the leaves.

HEENAN

Cardamine thalassica Heenan , spec. nov. ( Fig. 82 View FIGURE 82 )

Distinguished from C. glara by being entirely glabrous, with grey to grey-green leaves, leaflets with more distinct petiolules, inflorescence a raceme, shorter petals, and longer seeds.

Holotype:— NEW ZEALAND. Otago Land District, Hawkdun Range, Rambling Stream , bouldery colluvium, 7 December 2011, J. Barkla s.n. (CHR 619275!).

HEENAN NEW ZEALAND CARDAMINE REVISION

HEENAN

Perennial herb, single rosette or with lateral branches. Leaves up to 135 mm long, pinnatisect; lamina 17.0–65.0 × 16.0–42.0 mm, grey to grey-green, coriaceous, glabrous; petiole up to 92 mm long. Terminal pinna 5.7–14.0 × 4.5–9.5 mm, simple, broadly elliptic, elliptic-orbicular, ovate or broadly ovate; margin entire; lateral hydathodes absent; apex obtuse with a distinct hydathode; base obtuse. Lateral pinnae 2–6, 4.4–13.0 × 3.0– 9.5 mm, simple, broadly elliptic, elliptic-oribicular, ovate or broadly ovate; margin entire, lateral hydathodes absent; apex obtuse with a distinct hydathode; base obtuse; petiolule 1.8–10.5 mm long. Cauline leaves similar to rosette leaves but smaller, narrower, with fewer lateral pinnae, pinnae bases attenuate to cuneate; terminal pinna up to 20.0 × 5.0 mm, lateral pinnae up to 12.0 × 5.7 mm. Inflorescence racemose, sometimes with lateral racemes, each raceme 6–18- flowered; peduncle up to 250 mm long, 1.2–1.6 mm diam. at base, ascending, glabrous. Pedicels 10.0–22.0 mm long, 0.4–0.5 mm diam., glabrous. Sepals 2.1–2.4 × 0.8–1.4 mm, elliptic-oblong, saccate, green to red-brown, glabrous, margin white and membranous, apex obtuse, base truncate. Petals 3.6–4.5 × 1.2–1.4 mm, white, limb obovate; apex obtuse; base attenuate, tapering to a 1.0– 1.5 mm long claw. Stamens 6; median filaments 4, 2.4–3.1 mm long; lateral filaments 2, 2.1–2.9 mm long; anthers 0.3–0.4 mm long, cream, when dehiscent held slightly below the stigma. Ovary 2.5–3.2 mm long, 0.4–0.6 mm diam., ± terete, green-brown, glabrous; ovules 30–32; style c. 0.3 mm long, ± terete; stigma 0.4–0.5 mm diam. Siliques 21.0–34.0 × 1.1–1.3 mm, glabrous, style 0.7–1.0 mm long; valves green-brown at maturity and when dehiscent; replum 0.7–0.8 mm wide. Seeds 1.3–1.4 mm long, 0.8–0.9 mm wide, oblong to orbicular-oblong, yellow-brown; wing absent. FL Dec–Mar; FT Dec–Mar.

Representative specimens:— NEW ZEALAND. Otago. Nobbler Stream, Kakanui Mountains, 5 December

2006, J. Barkla s.n., CHR 591700; Mt Kakanui, 7 December 2013, D. Lyttle s.n., CHR 636076; St Bathans Range,

HEENAN NEW ZEALAND CARDAMINE REVISION

Phytotaxa 330 (1) © 2017 Magnolia Press • 141 10 January 2011, D. Lyttle s.n., CHR 636057. Cultivated. ex Mt Kakanui, Otago, cultivated experimental nursery, Landcare Research, 25 March 2014, P. B. Heenan s.n., CHR 636116; ex Mt Kakanui, Otago, cultivated experimental nursery, Landcare Research, 3 March 2016, P. B. Heenan s.n., CHR 588763.

Distribution and habitats:— New Zealand, endemic. Cardamine thalassica occurs in North Otago ( Fig. 83 View FIGURE 83 ), where it grows on dry, unstable bouldery colluvium and rocky slopes.

Conservation status:— Cardamine thalassica is assessed as having a conservation status of Threatened, Nationally Endangered (B1), with the qualifier Data Poor ( Townsend et al. 2008). The qualifier Data Poor is applied as additional information is required on the number and size of the populations and further field work is likely to discover new populations.

Etymology:—The specific epithet thalassica (Latin: greyish) alludes to grey to grey-green leaves.

Cardamine unguiculus Heenan , spec. nov. ( Figs. 13 View FIGURE 13 , 84 View FIGURE 84 )

Distinguished from C. polyodontes in having entirely glabrous leaves with 1–2 pairs of lateral leaflets, terminal and lateral leaflets with prominent marginal hydathodes and 2–7 shallow angular or weakly rounded lobes, narrower sepals, shorter and narrower petals, and seeds with a prominent apical wing.

Holotype:— NEW ZEALAND. The Haystack , Matiri Range, NW Nelson, 5000 ft, damp hollow on limestone ridge, March 1981, A. P. Druce s.n. (CHR 366093!).

Perennial herb, single rosette or with short lateral branches, stem and branches 1.0–3.0 mm diam. Leaves up to 115 mm long, pinnatisect; lamina 16.0–60.0 × 9.0–32.0 mm, green to green-brown, pinnae midrib and main veins green or red to red-brown on adaxial surface, membranous, glabrous, matt. Terminal pinna 5.5–30.0 × 8.0–29.0 mm, reniform, broadly reniform, to orbicular-reniform, margin with 1–7 shallow lobes and 1–7 conspicuous hydathodes, apex obtuse to truncate with a conspicuous hydathode, base weakly to strongly cordate. Lateral pinnae 2–4, 3.1–14.0 × 2.4–17.0 mm, reniform, orbicular-reniform, or broadly elliptic, margin with 1–5 shallow rounded lobes and 1–5 conspicuous hydathodes or entire, apex obtuse with a conspicuous hydathode, base cordate to obtuse, petiolule (0.6–)3.0– 5.3 mm long; petiole up to 70 mm long. Cauline leaves similar to rosette leaves but smaller, lateral pinnae absent or 2(–4); pinnae reniform, orbicular-reniform to elliptic, lobes shallowly rounded to narrowly triangular, hydathodes conspicuous. Inflorescence with 1–8 racemes, each raceme up to 17-flowered; peduncle up to 400 mm long, 1.0– 2.2 mm diam. at base, green or green-brown and mottled red-brown, spreading to ascending, glabrous. Pedicels 4.0–17.0 mm long, 0.4–1.0 mm diam., terete, glabrous. Sepals 2.0–3.2 × 0.7–1.3 mm, elliptic-oblong to broadly elliptic, ± saccate, green or green-brown, glabrous, margin white and membranous, apex obtuse, base truncate. Petals 3.0–6.0 × 1.5–2.7 mm, white, limb obovate to broadly elliptic; apex obtuse; base attenuate to cuneate, tapering to a 0.5–1.2 mm long claw. Stamens 6; median filaments 4, 2.5–4.1 mm long; lateral filaments 2, 1.8–3.3 mm long; anthers 0.4–0.8 mm long, cream to pale yellow, when dehiscent held at a similar height to or slightly below the stigma. Ovary 2.8–6.0 mm long, 0.5–1.0 mm diam., ± terete, green, glabrous; ovules 13–24; style 0.3–1.1 mm long, ± terete; stigma 0.3–0.5 mm diam. Siliques (8.5–)17.0–43.0 × 1.5–2.3 mm, glabrous, style 1.0–3.0 mm long; valves green to green-brown at maturity; replum 0.4–1.4 mm wide. Seeds 1.5–2.4 mm long, (0.6–)1.0– 1.5 mm wide, 0.4–0.6 mm thick, oblong to oblong-elliptic, green to green-brown; apex broad, obtuse, with prominent wing c. 0.1 mm wide present at apex. FL Nov–Mar; FT Nov–Aug.

Representative specimens:— NEW ZEALAND. Nelson. Loveridge Creek, Arthur Range, February 1991, A. P. Druce 1308, CHR 471900; Turks Head Range, January 1983, A. P. Druce s.n., CHR 393752; The Haystack, Matiri Range, March 1987, A. P. Druce s.n., CHR 366094; Glenroy Valley, March 1984, A. P. Druce s.n., CHR 401793; Kaihoka coast, November 1974, A. P. Druce s.n., CHR 277837; The Needle, Douglas Range, February 1985, A. P. Druce s.n., CHR 396002; Thorns Creek, April 1967, M. A. Ritchie & I. M. Ritchie s.n., CHR 534365/ CHR 534364; Spey River, January 1977, A. P. Druce s.n., CHR 310517; Karamea, 15 December 1984, D. K. Manning s.n., CHR 419070. Westland. Blue Bottle Creek, 4 December 1996, M. J. Newfield 29-96, CHR 511820/ CHR 511826; Blue Bottle Creek, 18 June 1997, R. P. Buxton s.n., CHR 512972; Blue Bottle Creek, 16 September 2009, P. B. Heenan s.n., CHR 617204; lower Kokatahi Valley, R. Mason & N. T. Moar 5180, 13 February 1958, CHR 97630; Kokatahi River valley, 9 January 1997, P. J. Bellingham 847, CHR 511347; Kokatahi River valley, 16 January 1997, P. J. Bellingham 862, CHR 511360; Waitahu River, 16 August 2002, D. A. Norton s.n., CHR 567732/CHR 567734; Blackwater Creek, 25 November 1923, W. Mackay s.n., CHR 60427; Wilberg Range, 27 April 1993, P. Wardle, R. P. Buxton & K. A. Ford s.n., CHR 500006. Southland. Diamond Creek, Milford Track,

HEENAN HEENAN NEW ZEALAND CARDAMINE REVISION

Phytotaxa 330 (1) © 2017 Magnolia Press • 143 23 July 1967, C. Tate s.n., WELTU 7487; Cascade River, Martyr Hill, 11 March 1978, Wardle, Lee & Johnson s.n., CHR 311270. Cultivated. ex Cedarmans Block, Lake Kaniere, Westland, cultivated experimental nursery, Landcare Research, 29 January 1997, P. B. Heenan 11/97, CHR 511039; ex Blue Bottle Creek, Westland, cultivated experimental nursery, Landcare Research, 20 April 1997, P. B. Heenan s.n., CHR 512611; ex Blue Bottle Creek, Westland, cultivated experimental nursery, Landcare Research, 16 September 2009, P. B. Heenan s.n., CHR 618356; ex Blue Bottle Creek, Westland, cultivated experimental nursery, Landcare Research, 16 September 2009, P. B. Heenan s.n., CHR 618352.

Distribution and habitats:— New Zealand, endemic. Cardamine unguiculus is known from several sites in north-west Nelson, Westland and western Southland (Fiordland) in the South Island ( Fig. 83 View FIGURE 83 ). It grows in a variety of habitats including a sphagnum swamp, on damp rock outcrops, and on limestone in damp hollows and tussockland.

Conservation status:— Cardamine unguiculus is assessed as having a conservation status of At Risk—Naturally Uncommon, with the qualifiers Data Poor and Sparse ( Townsend et al. 2008). The qualifier Sparse is applied as current knowledge infers C. unguiculus comprises mostly small and scattered populations. The qualifier Data Poor indicates additional information is required on the number and size of populations.

Etymology:—The specific epithet unguiculus (Latin: fingernail) alludes to the French manicure fingernail-like appearance of the apical wing on the seed.

Notes:—Leaf colour varies in C. unguiculus with this variation often occurring in the same geographic area, such as at Blue Bottle Creek, Lake Kaniere, Westland. At this site, some plants have leaves that are green with inconspicuous green veins and others green-brown with conspicuous red to red-brown veins.

Some of the Nelson collections of C. unguiculus were previously identified as belonging to the tagname entities C. “Tussock Race” (e.g., CHR 366094) or C. “Glossy Leaf” (e.g., CHR 419070); collections referrable to C. “Tussock Race” are herein named as C. polyodontes and those of C. “glossy leaf” named as C. chlorina .

Cardamine unicaulis Heenan View in CoL nom. nov. ≡ Cardamine hirsuta var. uniflora Hooker (1864: 12) View in CoL ≡ Cardamine heterophylla var. uniflora (Hook.f.) Cockayne (1909: 54) View in CoL ≡ Cardamine uniflora (Hook.f.) Allan (1961: 183) View in CoL , non Cardamine uniflora Michaux (1803: 29) View in CoL . Lectotype (fide Allan 1961, p. 184; Fig. 85 View FIGURE 85 ):— NEW ZEALAND. Colenso 1813 (lectotype K!). ( Fig. 8 View FIGURE 8 )

Typification notes:— The lectotype for C. hirsuta var. uniflora View in CoL is Colenso specimen number 1813 and this was sent by W. Colenso to J. D. Hooker with a letter dated 20 October 1848 (see St. George 2009, p. 236). Colenso’s correspondence stated for this entry: “1813. Cardamine View in CoL , minute species, shaded spots, cliffs nr. Porangahau.” Porangahau is located in southern Hawke’s Bay, North Island. The lectotype specimens feature the characteristic diminutive growth habit, small solitary flowers, and short silique .

Perennial herb, single rosette or with short lateral branches, stem and branches 0.9–2.5 mm diam. Leaves up to 85 mm long, simple or pinnatisect; lamina 1.8–25.0 × 1.3–14.0 mm, light green to green, membranous, glabrous to sparsely or moderately hairy on adaxial surface, margin and petiole, glabrous on abaxial surface. Terminal pinna 1.8–14.5 × 1.3–14.3 mm, simple, reniform, orbicular-rhomboid to broadly elliptic, apex obtuse with an inconspicuous hydathode, base truncate to cordate. Lateral pinnae absent or 1–4, 1.8–12.0 × 0.8–5.5 mm, orbicular, orbicular-rhomboid, to broadly elliptic, base often oblique, petiolule 0.2–1.7 mm long; petiole up to 60 mm long; hairs 0.2–0.4 mm long, spreading. Cauline leaves, usually simple, with fewer and narrower leaflets than rosette leaves, becoming smaller in all parts; upper leaves up to 10.0 × 1.6 mm, increasingly linear. Inflorescence with flowers solitary or corymbose, corymbs 2–6-flowered; peduncle up to 70 mm long, 0.4–0.6 mm diam. at base, spreading to ascending, glabrous. Pedicels 1.5–14.0 mm long, 0.2–0.4 mm diam., glabrous. Sepals 1.1–1.6 × 0.5–0.8 mm, elliptic-oblong to narrowly elliptic-oblong, saccate, green or red-brown, glabrous or sparsely hairy distally, margin white and membranous, apex obtuse, base truncate. Petals 3.5–4.7 × 1.7–2.5 mm, white, limb obovate; apex obtuse; base cuneate, tapering to claw up to c. 1.5 mm long. Stamens 6; median filaments 4, 1.0–2.0 mm long; lateral filaments 2, 0.8–1.7 mm long; anthers 0.4–0.5 mm long, cream to pale yellow, when dehiscent held at a similar height to or slightly below the stigma. Ovary 1.1–1.7 mm long, 0.15–0.25 mm diam., ± terete, green, glabrous; ovules 8–11; style 0.2–0.3 mm long, ± terete; stigma 0.15–0.25 mm diam. Siliques 8.0–13.5 × 0.9–1.2 mm, glabrous, style sessile or up to 1.0 mm long; valves green to green-brown at maturity; straw-coloured when dehiscent, replum 0.4–0.45 mm wide. Seeds 0.9–1.1 mm long, 0.6–0.8 mm wide, 0.25–0.45 mm thick, yellow-brown to henna; wing absent. FL Sep–Mar; FR Nov–Apr.

HEENAN

Representative specimens:— NEW ZEALAND. Wellington. Moawhango Gorge , December 1977, A. P. Druce s.n., CHR 323741; Manawatu Gorge, January 1944, A. P. Druce s.n., CHR 65055 . Otago. Earnscleugh , 4 April 2007, B. P. J. Molloy s.n., CHR 588296 . Southland. Bluff Hill, 4 January 1940, L. B. M[oore] s.n., CHR 23933; Centre Island, Foveaux Strait, March 1964, G. I. Collett s.n., CHR 151317; Haldane Bay, 5 February 1982, P. N. Johnson s.n., CHR 364558; Ocean Beach, 3 February 1957, M. E. Gillham s.n., CHR 111402. Stewart Island . Paterson Inlet, 18 November 1978, H. D. Wilson 789-94, CHR 369354; Little Hellfire Beach, 26 April 1978, H. D. Wilson 247, CHR 325114; Ocean Beach, Bluff, 9 February 1957, M. E. Gillham s.n., CHR 111439; Christmas Village , 18 February 1962, R. Melville 6401, CHR 145137. Cultivated. ex Red Rocks Stream, Wellington, cultivated Taita, Wellington, March 1967, A. P. Druce s.n., CHR 197169; ex Alexandra, Otago, cultivated Riccarton, Christchurch, Canterbury, 15 December 2006, B. P. J. Molloy s.n., CHR 588301; ex Alexandra, Otago, cultivated Riccarton, Christchurch, Canterbury, 19 January 2007, B. P. J. Molloy s.n., CHR 588302; ex Alexandra, Otago, cultivated experimental nursery, Landcare Research, Lincoln, 15 March 2010, P. B. Heenan s.n., CHR 618428; ex Maraetotara, Hawke’s Bay, cultivated Taita, Wellington, November 1962, A. P. Druce s.n., CHR 131960 .

Distribution and habitats:— New Zealand, endemic. Cardamine unicaulis is known from scattered localities throughout North Island, South Island and Stewart Island ( Fig. 83 View FIGURE 83 ). It mainly occurs in low altitude sites with the majority of collections being from coastal areas, but it does occur inland. It grows in a variety of habitats, including lake and pond margins, river banks, the base of rock outcrops, among coastal turf, sand and rocks and on limestone rocks. It seems to prefer partially shaded sites such as occur with an open forest canopy or among rock outcrops, and also moist areas such as muddy or damp ground.

Conservation status:— Cardamine unicaulis is assessed as having a conservation status of Data Deficient ( Townsend et al. 2008). Little is known about the number and size of the populations and what the threats are.

Etymology:—The specific epithet unicaulis (Latin: single stalk or stem) alludes to the mostly single flowering stems and retains the intent of the original species epithet uniflora (Latin: single flower).

Notes:—At least some of the plants (e.g., those cultivated from Red Rocks Stream, Wellington) included here

HEENAN NEW ZEALAND CARDAMINE REVISION

Phytotaxa 330 (1) © 2017 Magnolia Press • 145 in C. unicaulis were referred to as being aligned with the “mainland coastal race” by Pritchard (1957, pp. 82–83). Specimens from the Southland coast, Foveaux Strait, and Stewart Island have broader siliques and cauline leaves than the other collections.

Cardamine verna Heenan , spec. nov. ( Fig. 86 View FIGURE 86 )

Distinguished from C. alalata by its larger growth habit, matt leaves, terminal leaflet irregularly lobed, lateral leaflets with a longer petiolule, longer inflorescence, larger petals, and larger seeds.

Holotype:— NEW ZEALAND. Isolation Creek, Waima River, Marlborough Land District , 320 m, on gravels in river bed, 2 September 2009, P. B. Heenan s.n. (CHR 617208!).

Perennial herb, single rosette or with short lateral branches, stem and branches 0.9–2.0 mm diam. Leaves up to 120 mm long, pinnatisect; lamina 30.0–75.0 × 8.0–46.0 mm, green to brown-green, usually semi-coriaceous, glabrous. Terminal pinna 6.0–25.0 × 5.5–22.0 mm, usually simple, orbicular-rhomboid, rhomboid, broadly elliptic-rhomboid to broadly elliptic, often irregularly shaped, apex obtuse with a conspicuous hydathode; margin entire, irregularly 146 • Phytotaxa 330 (1) © 2017 Magnolia Press

HEENAN and shallowly lobed, with 2 distinct hydathodes; base obtuse to sometimes ± truncate. Lateral pinnae 2–8, 4.0–13.0 × 4.0–9.0 mm, orbicular-rhomboid, rhomboid, broadly elliptic-rhomboid to broadly elliptic, petiolule 2.0–15.0 mm long; petiole up to 60 mm long, glabrous or sparsely hairy toward base. Cauline leaves similar to rosette leaves, but with fewer and narrower leaflets, becoming smaller in all parts; upper leaves 5.0–10.0 × 2.0– 3.5 mm, increasingly linear, simple. Inflorescence racemose, usually with lateral racemes, each raceme 4–12-flowered; peduncle 100–350 mm long, 1.2–2.1 mm diam. at base, spreading to ascending, glabrous. Pedicels 4.0–22.0 mm long, 0.3–0.6 mm diam., glabrous. Sepals 2.4–2.6 × 1.2–1.4 mm, elliptic-oblong to broadly elliptic, saccate, green or red-brown, glabrous, margin white and membranous, apex obtuse, base truncate. Petals 5.0–11.5 × 2.5–5.5 mm, white, limb obovate to broadly elliptic-obovate; apex obtuse; base cuneate to obtuse, tapering to a 1.0–2.0 mm long claw. Stamens 6; median filaments 4, 2.5–4.2 mm long; lateral filaments 2, 2.4–3.5 mm long; anthers 0.7–0.8 mm long, cream to pale yellow, when dehiscent held slightly at a similar height to or slightly below the stigma. Ovary 4.2–4.5 mm long, 0.5–0.7 mm diam., ± terete, green, glabrous; ovules 19–22; style 0.5–1.3 mm long, ± terete; stigma 0.7–0.8 mm diam. Siliques 17.0–40.0 × 1.2–1.5 mm, glabrous, style 0.2–1.4 mm long; valves green at maturity, straw-coloured when dehiscent; replum 0.5–0.8 mm wide. Seeds 1.2–2.1 mm long, 0.8–1.1 mm wide, 0.3–0.4 mm thick, orbicular-oblong to broadly oblong, yellow-brown to henna; wing present at apex and base, especially well-developed at apex, 0.1–0.2 mm wide. FL Aug–Dec; FT Nov–Apr.

Representative specimens:— NEW ZEALAND. Marlborough. Mt Ben More, November 1973, A. P. Druce s.n., CHR 249263–249265; Mt Ben More, November 1973, A. P. Druce s.n., CHR 249191; Chalk Range, 24 December 1953, R. Mason & D. R. McQueen 2634, CHR 85089; Isolation Creek , December 1975, A. P. Druce s.n., CHR 313029; Isolation Creek , April 1981, A. P. Druce s.n., CHR 387770; Ure Valley , G. Simpson s.n., CHR 18750; Isolation Creek , 2 September 2009, P. B. Heenan s.n., CHR 617194; Woodside Creek , November 1975, A. P. Druce s.n., CHR 313032. Cultivated. ex Waima River , Marlborough, cultivated Pinehaven, Hutt Valley , Wellington, February 1985, A. P. Druce s.n., CHR 394349; ex Waima River , Marlborough, cultivated Pinehaven, Hutt Valley , Wellington, October 1985, A. P. Druce s.n., CHR 394311; ex Isolation Creek , Marlborough, cultivated experimental nursery, Landcare Research, 3 December 2009, P. B. Heenan s.n., CHR 618346–618349; ex Isolation Creek , Marlborough, cultivated experimental nursery, Landcare Research, 19 October 2009, P. B. Heenan s.n., CHR 618406 .

Distribution and habitats:— New Zealand, endemic. Cardamine verna is known from limestone in the vicinity of Ben More and Isolated Hill, in a restricted geographic area between the northern end of the Chalk Range and lower parts of the Waima River, Marlborough (South Island) ( Fig. 83 View FIGURE 83 ). It mainly occurs in river and stream beds among rocks and stones, and on alluvium and rock outcrops.

Conservation status:— Cardamine verna is assessed as having a conservation status of Threatened, Nationally Vulnerable (B1), with the qualifier Data Poor ( Townsend et al. 2008). The qualifier Data Poor is applied as additional information is required on the number and size of the populations.

Etymology:—The specific epithet verna (Latin: spring) refers to the early flowering of this species, beginning in August and September.

Notes:—There are two flower size morphs in C. verna ( Fig. 86F View FIGURE 86 ). The larger flower is up to 13 mm diameter and has petals 7.5–11.0 mm long, median filaments 3.2–4.2 mm long, stigma 0.8–1.0 mm diam., and the stigma protrudes above the dehiscent anthers. The smaller flowered morph has 9–10 mm diameter flowers, petals 5.0–7.0 mm long, median filaments up to 2.7–2.9 mm long, stigma 0.6–0.7 mm diam., and the stigma is placed below the dehiscent anthers. The larger flower and exserted stigma infers the large-flowered morph may be adapted to outcrossing, whereas the small-flowered morph with the stigma held below the anthers indicates it may be suited to self-pollinating.

Previously known by the tagnames Cardamine “Waima” or “Benmore”.

Hybrids:— Cardamine verna × C. corymbosa . A single plant has features that indicate it maybe 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

HEENAN NEW ZEALAND CARDAMINE REVISION

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).

Incertae sedis

The following names were published by Schulz (1903). Type specimens for Schulz names were deposited in B, but this herbarium was severely damaged by fire in 1943 and many types were destroyed. No types have been located for the taxa listed below (R. Vogt, pers. comm., 18 October 2013). The descriptions for these taxa are inadequate to allow for the selection of a neotype and they are therefore treated as incertae sedis. Since these Schultz (1903) names are at infraspecific rank they will not be in conflict with any of the names proposed in this revision.

Cardamine heterophylla View in CoL prol. micrantha O.E. Schulz (1903: 488)

Cardamine heterophylla View in CoL prol. macrantha O.E. Schulz (1903: 489)

Cardamine heterophylla View in CoL prol. macrantha var. leiocarpa O.E. Schulz (1903: 489)

Cardamine heterophylla View in CoL prol. macrantha var. hirtella O.E. Schulz (1903: 489)

Cardamine heterophylla View in CoL prol. macrantha var. hirtella var. macrostylis O.E. Schulz (1903: 489)

Indeterminate specimens

A number of specimens were unable to be assigned to a species accepted in this revision and have been treated as indeterminate. Many of the specimens in this category lack relevant diagnostic characters as they were either poor quality and/or sterile; it is unlikely these specimens will be able to be assigned an appropriate species name. Another group of specimens could not be placed in a named species but either had some unique character or could be grouped together by their general similarity and/or geographical proximity. These groupings possibly include unnamed novelties and are discussed below. They require further research, including field survey and cultivation experiments.

Cardamine aff. alalata ( Fig. 87 View FIGURE 87 )

Vouchers:— Southland. Princess Range, c. 5000 ft, 26 January 1924, J. Spedon s.n., WELTU 20223; Lake Monk, 19 January 1960, M. J. A. Simpson 1843, CHR 110802; Lake Monk, 19 January 1960, M. J. A. Simpson 1843, CHR 110806.

Distribution:— Known from several collections in Fiordland (western Southland).

Recognition:— These plants are similar to C. alalata in having a compact growth habit and probably rhizomatous stems, pinnatifid leaves with prominent hydathodes and a compact inflorescence, but they differ in lacking the wing on the seed. Their compact growth, pinnatifid leaves and the short and compact inflorescence is similar to C. latior , but the siliques are more slender and seeds smaller than that species.

Cardamine “Eweburn”

Voucher:— Otago. Eweburn Dam, W. of Naseby , 770 m, 18 December 1976, T. Partridge & J. Child s.n., OTA 36555 About OTA .

Distribution:— Known from a single collection comprising five specimens from Eweburn Dam, near Naseby, north Otago.

HEENAN

Recognition:— Recognised by its simple or 2-lobed, sparsely hairy, 10–20 mm long leaves, inflorescence sparsely hairy, and densely hairy siliques. The hairs on the silique are spreading to weakly retrorse but their density varies, with the replum sparsely hairy and valves densely hairy. Among the New Zealand species the presence of

HEENAN NEW ZEALAND CARDAMINE REVISION

hairs on the replum is unique to these specimens. These specimens are most similar to C. intonsa in having the hairy silique valves, and further research may result in them being assigned to that species.

Cardamine “Fiordland” ( Fig. 87 View FIGURE 87 )

Vouchers:— Southland. Marrington Peaks, 9 January 1957, M. J. A. Simpson s.n., CHR 94174; Caroline Burn, Lake Hauroko, 24 November 1975, C. Sutcliffe, M. Craighead & Z. Williams s.n., CHR 307647; camp to Irene Saddle, 26 June 1957, M. Cookson s.n., CHR 96293; Dove Pass, 4 February 1982, J. V. Morrison s.n., CHR 594188; Mount George, 21–25 March 1977, P. Garnock-Jones, W. Lee, J. Anderson & D. Given 10,373, CHR 306262; Lake Wapiti, 23 February 2016, R. Hindmarsh-Walls s.n., CHR 588999.

Distribution:— Known from several collections in Fiordland (western Southland).

Recognition:— Recognised by its pinnatisect leaves with the terminal lobe usually distinctly trilobed, prominent leaf margin hydathodes, one pair of lateral leaflets, and corymbose inflorescence with long pedicels.

Cardamine “northern robust”

Vouchers:— North Auckland. Hingaia Point, Little Barrier Island, 28 August 1997, P. B. Heenan s.n., CHR 513346, Te Maraeroa Flat, 29 November 1978, A. E. Esler & R. E. Beever 78113, CHR 324839; Te Titoki Point, Little Barrier Island, 13 June 1984, R. Bieleski & R. Beever 84044, CHR 411953; Little Barrier Island, July 1952, J. E. Moore s.n., CHR 78464; Motukino (Fanal) Island, Mokohinau Islands, 15 September 1994, E. K. Cameron 7731 & P. J. de Lange, AK 226078/CHR 486773; Mokohinau Islands, August 1965, G. I. Collett s.n., CHR 183340; Whatapuke Island, 27 October 1968, M. A. & I. M. Ritchie s.n., CHR 186850; Kaawa Creek, Raglan Co., 20 November 1958, R. Mason & N. T. Moar 6095, CHR 109197; Otorohonga, Raglan Co., 1 December 1958, R. Mason & N. T. Moar 6601, CHR 109929. GISBORNE: Waiotahi Domain, Opotiki, December 1976, M. Heginbotham s.n., CHR 370288.

Distribution:— Known from several collections in North Auckland and several offshore islands, and a single collection from Gisborne.

Recognition:— Densely leafy plants, with robust with leaves upto 100 mm long with a large terminal lobe up to 25 × 27 mm, 2–4 large lateral lobes, and the leaves persistent in fruit. The cauline leaves are well-developed with prominent terminal and lateral lobes.

Cardamine “Turoa” ( Fig. 87 View FIGURE 87 )

Voucher:— Wellington. Mangaturuturu River, Mt Ruapehu, 12 March 2015, P. B. Heenan s.n., CHR 640349. Cultivated. ex Turoa Alpine Village, near waterfall, collected P. J. de Lange, cultivated experimental nursery, Landcare Research, 6 March 2014, P. B. Heenan s.n., CHR 636106.

Distribution:— Known from near Turoa Alpine Village, northern Wellington, Mt Ruapehu, central North Island.

Recognition:— Recognised by pinnatisect hairy leaves with obvious but not protruding marginal hydathodes, decumbent and sprawling inflorescences up to 50 cm long, and inflorescence with leafy rosettes scattered along its length. The inflorescence rosettes are unusal and have not been recorded in any other species.

Cardamine “west Otago ” ( Fig. 87 View FIGURE 87 )

Vouchers:— Otago. Staircase Creek, Hector Mountains, 1 February 2007, M. Thorsen 023/07, CHR 591775; Harris Mountains, 25 February 2009, M. Thorsen 082/09, CHR 605870.

Cultivated. ex Harris Mountains, Otago, cultivated experimental nursery, Landcare Research, 3 December 2009, P. B. Heenan s.n., CHR 636105; ex Harris Mountains , Otago, cultivated experimental nursery, Landcare Research, 3 December 2009, P. B. Heenan s.n., CHR 618341; ex Harris Mountains , Otago, cultivated experimental nursery, Landcare Research, 2 December 2009, P. B. Heenan s.n., CHR 618330 .

Distribution:— Known from the Hector Mountains and Harris Mountains in western Otago. A number of other collections maybe conspecific that come from western Otago (e.g., Forbes Mountains, A. F Mark s.n., OTA 22320; Emily Pass, Routeburn Valley, A. F. Mark & M. L. Burke s.n., OTA 21022; Fiery Col, Humbolt Mountains, A. F. Mark s.n., OTA 22350; Deep Creek, Harris Mountains, J. Barkla s.n., CHR 572422) and Southland (e.g., Homer Tunnel, Fiordland, L. C. Bliss s.n., OTA 12116).

Recognition:— Recognised by its coriaceous green pinnatisect leaves, petiolule absent or short, inflorescence with siliquas erratically placed and less than 15 mm long.

HEENAN

Cardamine “Ultra”

Voucher:— Marlborough. Red Hills , December 1980, A. P. Druce s.n., CHR 387497 .

Distribution:— Known from one collection on ultramafic rocks where it is said to be common. Cardamine dactyloides also occurs on ultramafic rocks in the same area.

Recognition:— Recognised by its angular leaves with prominent hydathodes, 1–2 pairs of lateral pinnae, and oblong orange-brown seeds.