A L. oleraceo caulibus gracilibus flexilibus, foliis pallide viridibus membranaceis, lanceolatis elliptico-lanceolatis vel anguste ellipticis, apice prominente acuto vel subacuto, marginibus serratis dentibus prominentibus profundis, et sequentia nucleotidorum DNA distinguenda.

(Fig. 9). New Zealand: Canterbury, Lincoln, Landcare Research experimental nursery, ex Banks Peninsula, rock stack near Island Bay, May 2011, P. B. Heenan s.n., CHR 616211!

The specific epithet ‘aegrum’ is derived from Latin, meaning ‘diseased and/or sick body’ and refers to the condition of the plants in the single known extant wild population on the unnamed rock stack near Island Bay, many of which are infected with turnip mosiac virus (Fletcher et al. 2009).

(Figs 10–14). Tap-rooted, strongly pungent smelling, perennial herb. Growth habit open, up to 50 cm tall, stems arising from basal woody stems. Stems upright, slender, flexible; mature stems woody, 100–500 × 10–12 mm, often devoid of foliage on middle and lower parts of stems; new stems 80–200 × 3–4 mm, leafy, glabrous. Leaves glabrous, membranous, light green, planar, pellucid glands sometimes scattered on abaxial surface, rosette and stem leaves usually withering, variable in size and shape. Leaves of young and vigorous plants and stems: lamina 35–80 × 10–20 mm, lanceolate, elliptic-lanceolate, narrowly elliptic; apex subacute, with a single prominent tooth; margin singly serrate, with 8–12 pairs of teeth; teeth up to 3.5 mm deep, not overlapping; base attenuate, tapering to distinct petiole. Leaves of mature plants and cauline stems: lamina 30–65 × 4–11 mm, narrowly lanceolate, elliptic-lanceolate, narrowly elliptic; apex acute to subacute, with a single prominent tooth; margin singly serrate in upper and/or lower half, with 2–10 pairs of teeth; teeth up to 1.2 mm deep, not overlapping; base attenuate, tapering to petiole. Inflorescence terminal and lateral, racemose, often branched, up to 95 mm long, rachis up to 2.1 mm diam., glabrous; pedicels 4–10 mm long, erecto-patent, glabrous. Flowers 4.0–4.5 mm diam. Sepals 4, 1.0–1.5 mm long, saccate, green, apex obtuse, margin white, shape dimorphic; lateral sepals broad, 1.0–1.1 mm diam., orbicular, abaxial surface often hairy, hairs entirely eglandular or with glandular tip, 0.2–0.3 mm long; median sepals narrow, 0.8–0.9 mm diam., broadly elliptic, glabrous. Petals white, 2.3–2.5 × 1.2–1.4 mm, spreading, claw 0.7–0.9 mm long; limb broadly elliptic to orbicular, apex obtuse to rounded. Stamens 4; filaments 1.7–2.0 mm long, base 0.3–0.4 mm diam., equal; anthers 0.4–0.5 mm long. Ovary 0.9–1.0 × 0.9–1.0 mm, broadly ovate to broadly elliptic, green, apex usually with shoulders; style 0.3–0.4 mm long, cylindrical; stigma 0.3–0.4 mm diam. Nectaries 4, 0.2–0.3 × c. 0.1 mm, oblong, green. Silicles cartilaginous when fresh, coriaceous when dry, 4.0–4.7 × 3.2–3.5 mm, elliptic-rhomboid to orbicular-rhomboid, apex shallowly notched, valves pale brown, glabrous, not winged; style 0.3–0.4 mm long, exserted. Seeds 1.6–1.7 × 0.9–1.0 mm, narrowly ovoid, brown to orange-brown, not winged. FL Mar. FR Mar.

New Zealand (South Island): Banks Peninsula, Waikerikeri, n.d., R. M. Laing s.n., (AK 4463); Banks Peninsula, Akaroa, Akaroa Head, Island Bay, Unnamed Islet, July 2002, N. Head s.n., AK 283510. Cultivated (New Zealand): Landcare Research experimental nursery, Lincoln, ex Akaroa, Akaroa Head, Island Bay, Unnamed Islet, 5 September 2008, P. J. de Lange 7363 & G. Houliston, (AK 303515);Landcare Research experimental nursery, Lincoln, ex Island Bay, Banks Peninsula, 13 January 2010, P. B. Heenan s.n., (CHR 609820); Landcare Research experimental nursery, Lincoln, ex Island Bay, Banks Peninsula, 13 January 2010, P. B. Heenan s.n., (CHR 609821); Landcare Research experimental nursery, Lincoln, ex Island Bay, Banks Peninsula, 1 April 2009, P. B. Heenan s.n., (CHR 609804); Landcare Research experimental nursery, Lincoln, ex Island Bay, Banks Peninsula, 11 August 2009, P. B. Heenan s.n., (CHR 609792).

Distribution (Fig. 15). Endemic. New Zealand, South Island, Canterbury, Banks Peninsula, Island Bay, unnamed rock stack.

Known from a single wild population on a small and unnamed island near Island Bay. Here it grows in open and disturbed areas among petrel burrows and Disphyma australe (W.I.Aiton) N.E.Br. subsp. australe (Fig. 6). Additional populations of Lepidium aegrum have been established in Canterbury on Motunau Island (Pegasus Bay), Quail Island (Lyttleton Harbour), and at Stony Bay (Banks Peninsula).

Recognition. Lepidium aegrum is distinguished by its slender flexible stems (Figs 11, 12), light green membranous leaves that are lanceolate, elliptic-lanceolate or narrowly elliptic with a prominent acute to subacute apex, and margins that are serrate with prominent and deep teeth (Fig. 13).

Conservation Status. Lepidium aegrum is known from a single wild population and three additional populations that have been established with nursery-raised plants. The single wild population and unknown recruitment patterns means that this species is especially vulnerable to stochastic events. Furthermore, turnip mosiac virus, which deforms and retards growth, is common in plants at the wild population, and this will likely reduce reproductive success (Fletcher et al. 2009). Using the New Zealand Threat Classification System (Townsend et al. 2008), Lepidium aegrum qualifies as Threatened/Nationally Critical. We recommend appending the qualifiers ‘CD’ (Conservation Dependent – as it has been established and is being managed at Motunau and Quail islands, and at Stony Bay), ‘DP’ (Data Poor – to reflect uncertainty over plant numbers and population trends), ‘OL’ (One location – since it is known from a single wild population).

A L. obtusato habitu suberecto parce ramoso et sine rhizome, caulibus arcutis potius quam flexuosis, foliis rosulae et caulis inferi profunde denticutis persistente oblanceolato-spathulatis ad obovato-spathulatis, foliis caulis superi sparsis lanceolatis ad anguste oblanceolatis vel obdetoideis truncatis et profunde dentatis (raro integris) paribus dentorum usque ad 6 profunde incises et sparsim foliaceis, racemis longioribus quibus non obscuris caulibus vegetativis siliclisque parvioribus et angustioribus differt.

New Zealand (Fig. 16): Cliffs between Karekare and Manukau Heads, January 1917, T. F. Cheeseman s.n., AK 4474! Isotypes: AK 206570!, WELT SP030095!

The epithet ‘amissum’ is derived from the Latin ‘amissus’ meaning ‘lost’ and is used here to refer to the loss though extinction, of this plant from the New Zealand flora.

(Figs 17–20). Glabrous, suberect, sparingly branched, succulent, perennial, herb forming open patches up to c. 1 m diam. Rootstock stout, woody, exposed portion rough, covered in old dead stem and rosette-leaf remnants admixed with actively growing stems. Stems arching, widely spreading, possibly with apices weakly ascendent, succulent, mature stems woody, 200–300 × 3–6 mm, densely leafy near base, leaves more widely spaced along upper stems. Leaves glabrous, coriaceous, probably succulent, dark green, planar, variable size and shape. Rosette leaves persistent at fruiting 71.7–95.2 × 12.4–23.2 mm, oblanceolate-spathulate to obovate-spathulate; apex, truncate, praemorse, with 3–8 deeply incised teeth; margin coarsely and often irregularly incised or dentate, often weakly bidentate, with 24–36 pairs of teeth; teeth up to 4.8 mm deep, irregular in size, protruding beyond leaf outline; base narrowly attenuate to cuneate, ± decurrent, petiole distinct, 43.4–55.2 × 1.4–2.3 mm long, slightly winged, or not, channelled. Lower stem leaves similar to rosette leaves, apparently persistent, widely spaced, gradually decreasing in size toward inflorescence; petioles distinct, slightly winged or not. Upper stem leaves much reduced; lamina 10.3–27.5 × 3.4–7.6 mm, lanceolate, narrowly oblanceolate, lanceolate to narrowly obdeltoid, apex truncate, with 3–4 prominent teeth, margins prominently toothed in upper ⅓ of lamina with 4–6 deeply incised pairs of teeth (rarely entire except for apex), base cuneate to broadly cuneate; petiole distinct or indistinct, up to 6 mm long when present, channelled. Inflorescence racemose, terminal and lateral, conspicuous, sparingly leafy and unfettered by associated vegetative leafy stems; racemes 22–68 mm long, rachis 0.9–1.6 mm diam., glabrous; pedicels 2.8–3.2 mm long at flowering, erecto-patent, elongating somewhat after anthesis, glabrous. Flowers 4.3–4.6 mm diam. Sepals 4, saccate, overlapping at base, green, apex obtuse, margin white, shape and size dimorphic; lateral sepals 2.2–2.9 × 2.1–3.0 mm, suborbicular, mostly glabrous, sometimes sparsely hairy, hairs 0.2–0.4 mm long, caducous; median sepals 1.9–2.9 × 1.5–1.7 mm, broadly elliptic to obovate, abaxial surface glabrescent, sparsely hairy, hairs 0.2–0.4 mm long, caducous. Petals white, 1.3–1.8 × 1.3–1.8 mm, erect, claw minute, 0.2–0.3 mm; limb orbicular, apex obtuse. Stamens 4, ± equal lengths, 1.2–1.8 mm long, base 0.6–1.0 mm wide; anthers 0.6–0.8 mm long, yellow, pollen yellow. Ovary 1.3–1.8 × 1.3–1.7 mm, broadly ovate to suborbicular green, apex distinctly notched; style 0.3–0.5 mm long, cylindrical below, spreading at apex; stigma 0.5–0.6 mm diam. Nectaries 4, green, 0.12–0.14 × c. 0.09 mm, narrow oblong, apex obtuse. Silicles 3.4–4.5 × 2.9–3.9 mm, broadly ovate, oval to obovate, apex prominently notched, valves yellow-green (in dried specimens), glabrous, slightly winged; style 0.2–0.5 mm long, not or only slightly exserted. Seeds 2.0–2.7 × 1.8–2.0 mm, obovate, broadly obovate, brown to orange-brown, not winged. FL: Dec–Jan. FR: Dec–Jan.

New Zealand (North Island): North Auckland, Cliffs north of the Manukau heads, December 1870, T. F. Cheeseman s.n., (AK 4473).

(Fig. 15). Endemic. New Zealand, North Island, where it was recorded from ‘between Manukau Heads and Karekare’ (Cheeseman 1906, 1925), not the ‘Titirangi’ noted by Kirk (1899). Lepidium amissum was last collected from the west Auckland coastline in 1917 and is presumed to be extinct.

Lepidium amissum had previously been included in Lepidium obtusatum, perhaps because the species have no obvious floral differences and both have deeply notched and winged silicles with tapering bases. However, from Lepidium obtusatum, Lepidium amissum differs by the non-rhizomatous, suberect growth habit; arching, sparingly leafy stems (Fig. 16); long persistent, deeply, and sharply incised rosette and lower stem leaves (Figs 17, 18); upper stem leaves which are lanceolate, narrowly oblanceolate, to narrowly obdeltoid and mostly prominently toothed, with up to 6 pairs of deeply incised teeth, and with a truncate apex bearing 3–4 prominent teeth. The inflorescences of Lepidium amissum are larger, only sparsely leafy when mature, and without associated vegetative stems (Fig. 19). The silicles are smaller than those of Lepidium obtusatum (Fig. 20).

Little is known about the habitat preferences and ecology of Lepidium amissum, beyond that it grew on sea cliffs (Cheeseman 1906, 1925).

Lepidium amissum is considered to be extinct. There have been repeated unsuccessful surveys for this species over the last fifty years by various botanists. Aside from the type the species is known from only one collection made in 1917. This collection and the type came from an ill defined area of coastline stretching for some 6 km from the Manukau Heads to Karekare. Today this coastline is protected within the Centennial Park, Waitakere Ranges. However, during the period when this species was gathered, all of the coastline was being modified as a result of kauri (Agathis australis (D.Don) Lindl.) logging (see Harvey 2001, 2006; MacDonald and Kerr 2009), and this may have contributed to its eventual extirpation. It does seem that Lepidium amissum was already uncommon at the time of its discovery and it is possible that the gatherings made by Thomas Cheeseman where sufficient to cause its extinction as they are copious and comprise many stems.

“SOUTH Island: Queen Charlotte Sound and Astrolabe Habrour, A. Richard, l.c.”

(Fig. 21, designated here): Herbarium Richard, Paris!

Although Kirk (1899) did not explain his choice of epithet, ‘banksii’ commemorates Sir Joseph Banks FLS, FRS (1743–1820) who together with Dr Daniel Solander made the first gatherings of New Zealand plants to be described by European botanists during the Endeavour voyage of discovery (1768–1771).

= Lepidium banksii var. ovatum Kirk, Stud. Fl. N. Z., 35 (1899)

(fide Allan 1961): Northwest Bay, Pelorus Sound, J. Rutland s.n., n.d., WELT SP079012!

Although Kirk (1899) did not explain his choice of epithet, the protologue makes it clear that “ovatum” was chosen to refer to the shape of the mature silicles.

The key provided by Kirk (1899, p. 34) used the name “Lepidium forsteri” instead of Lepidium banksii. This is considered to be a nomen nudum.

(Figs 22–25). Tap-rooted, strongly pungent smelling, perennial herb. Growth habit dense, stems closely placed, 20–50 cm tall. Stems upright to spreading, stout, barely flexuous; mature stems woody, 100–500 × 3–8 mm, often devoid of foliage on middle and lower parts of stems. Leaves glabrous, coriaceous, green, planar, rosette and stem leaves usually withering, variable in size and shape. Leaves of young and vigorous plants and stems: lamina 20–40 × 6–15 mm, oblanceolate-spathulate, obovate; apex obtuse, often with up to 3 or 4 teeth; margin coarsely serrate, with 15–21 pairs of teeth; teeth up to 2.0 mm deep, irregular in size, protruding beyond leaf outline; base attenuate to cuneate, petiole distinct; petiole up to 35.0 × 1.3–2.8 mm, channelled. Leaves of mature plants and cauline stems: lamina 8–25 × 3–6 mm, linear-oblanceolate, obovate; apex obtuse to truncate, often with up to 3 or 4 teeth; margin serrate in upper half, up to 7 pairs of teeth; not overlapping, up to 1.5 mm deep, often protruding beyond leaf outline; base attenuate to cuneate, usually tapering to ± distinct petiole, sometimes appearing sessile; petiole up to 8.0 × 1.0–1.8 mm, channelled. Inflorescences terminal and lateral, racemose, 20–80 mm long, rachis 0.6–1.4 mm diam., glabrous or sometimes with pale clavate hairs; pedicels 5–8 mm long, erecto-patent, with pale clavate hairs on adaxial surface, hairs 0.1–0.15 mm long. Flowers 4.0–4.5 mm diam. Sepals 4, saccate, overlapping at base, green, apex obtuse, margin white, shape and size dimorphic; lateral sepals 1.6–2.1 × 1.1–1.5 mm, orbicular, glabrous; median sepals 1.5–1.9 × 0.9–1.1 mm, broadly elliptic, abaxial surface glabrous or sparsely hairy, hairs 0.2–0.4 mm long. Petals white, 1.8–2.0 × 0.1–0.9 mm, erect, claw indistinct; limb narrrowly obovate, elliptic or filiform, often irregular in shape, apex obtuse to subacute. Stamens 4, ± equal lengths, 1.2–1.7 mm long, base 0.6–0.9 mm wide; anthers 0.4–0.7 mm long, yellow or sometimes violet. Ovary 1.4–1.6 × 1.0–1.6 mm, broadly ovate, green, apex round or sometimes weakly shouldered; style 0.2–0.3 mm long, cylindrical below, spreading at apex; stigma 0.45–0.5 mm diam. Nectaries 4, 0.2–0.4 × c. 0.1 mm, oblong, green. Silicles cartilaginous when fresh, coriaceous when dry, 4.5–5.5 × 4.0–5.0 mm, broadly ovate, apex notched, base cordate, valves green maturing yellow-green, glabrous, slightly winged; style 0.2–0.3 mm long, exserted. Seeds 1.8–2.3 × 1.0–1.2 mm, obovate or obovate-elliptic, brown to orange-brown, not winged. FL Nov–Jan. FR Nov–Jan.

New Zealand (South Island): Tennyson Inlet, Tawa Bay, January 1910, E. Phillips-Turner 93, (AK 100087); Kenepuru Head, Pipi’s Beach, 1917, J. H. MacMahon s.n., (WELT SP030112); Kenepuru, n.d., n.c., , (WELT SP030113); ?Queen Charlotte Sound, n.d., J. H. MacMahon s.n., (WELT SP081929); Boulder Bank, November 1908, F. G. Gibbs s.n., (CHR 81737); Abel Tasman National Park, Totaranui Headland, December 1961, A. E. Esler s.n., (AK 218197); Abel Tasman National Park, Totaranui Headland, 27 January 1963, A. E. Esler s.n., (CHR 181143); Abel Tasman National Park, Totaranui Point, 12 January 1984, D. R. Given 13871 & A. D. Given, (CHR 509034); Abel Tasman National Park, Totaranui, 12 January 1986, D. R. Given 14106, (CHR 420352); Abel Tasman National Park, Totaranui Trig, 9 January 1984, D. R. Given 13505 & B. A. Duncan, (CHR 416178); Abel Tasman National Park, Separation Point, 28 January 1997, P. J. de Lange 3249 & G. M. Crowcroft, (AK 231008); Karamea, 28 January 1985, P. Wardle s.n., (CHR 446772); Bird Island, Waimea Estuary, 1946, C. Baas s.n., (CHR 278958). Cultivated (New Zealand). Landcare Research experimental nursery, Lincoln, ex Totaranui, Nelson, 17 November 2005, P. B. Heenan s.n., (CHR 504666); Landcare Research experimental nursery, Lincoln, ex Abel Tasman National Park, 6 August 2006, P. B. Heenan s.n., (CHR 609830).

(Fig. 15). Endemic. New Zealand, South Island, Marlborough Sounds (Queen Charlotte, Kenepuru and Pelorus Sounds), Nelson (Waimea Estuary, Totaranui, Separation Point, Wainui Inlet), north-west Nelson (Karamea).

Lepidium banksii is recognised by the clavate hairs on the pedicels (Fig. 24A), mostly filiform petals, styles that spread at the apex into a broad plate (Fig. 24B), and silicles that have a prominent apical notch (Fig. 25). In comparison, the styles of the other Lepidium species except for Lepidium seditiosum, are cylindrical for their whole length.

Lepidium banksii is a strictly coastal species favouring rubble slopes, boulder beaches, exposed rocky headlands (Fig. 22), and sparsely vegetated cliff faces usually near penguin colonies or low lying, estuarine shell banks and sand and shell barrier islands used as high tide roosts by wading birds.

Lepidium banksii is currently listed as ‘Threatened/Nationally Critical _{CD, EF}’ by de Lange et al. (2009). Both de Lange (2006) and Courtney (2009) describe the conservation management of this species, noting that there are probably no naturally occurring plants of Lepidium banksii left in the wild, with all those known the result of deliberate plantings and sowing of seed. As such, the qualifier ‘EW’ (Extinct in Wild) should be added to the current threat listing. The most recent census of the numbers of Lepidium banksii present in the ‘wild’ records 113 plants (Courtney 2009). Survival of this species is now completely dependent on direct hands-on management (de Lange 2006; Courtney 2009).

A Lepidio oleraceo sympatrico sed foliis in caule medio ad supero anguste lanceolatis ad linearo-lanceolatis dentibus acicularibus, apicibus foliorum decresento-acuminatis, et pedicellis minutis et sparsim puberuli differt, et sequentia nucleotidorum DNA distinguenda.

Kermadec Islands (Fig. 26): Macauley Island, north-east coast, 3 July 2006, J. W. Barkla M29 & T. C. Greene, AK 297694! Isotype: CHR 552474!

The epithet ‘castellanum’ from the Latin ‘associated with a fort, fortress or castle’ refers to the remote, craggy, castle-like habitats of this species in the southern Kermadec Islands group on which islands it is endemic.

(Figs 27–32). Tap-rooted, strongly pungent smelling, much-branched, leafy perennial shrub, up to 1.8 × 2.0 m. Tap-root up to 800 mm long, ± napiform and/or scarcely branched. Rootstock 6–10 mm diam., woody, exposed portion smooth. Stems persistent, arising from rootstock base and basal portion of main central stem, closely packed, woody, erect, weakly angled to ± terete, glabrous; mature stems 3.8–8.2 mm diam., 0.3–1.8 m long; red-green to yellow-green, brittle, bases bearing numerous leaf abscission scars otherwise mostly leafy from mid-stem to apex at flowering; middle stems dark green to red-green, fleshy and pliant, initially ± square, prominently angled, becoming ± terete with age. Leaves coriaceous, fleshy, green to dark green, rosette-leaves absent, stem leaves withering with age; basal stem leaves 88.5–120.0 × 14.5–30.0 mm, lamina broadly lanceolate to lanceolate, margin ± deeply and ± evenly incised, teeth in 50–90 ± equal pairs, 0.5–2.9 mm long, protruding beyond leaf line, narrowly deltoid, to deltoid, leaf apex truncate-praemorse often deeply lacerate, teeth 3–5, cut 2.4–4.2 mm to lamina, narrowly deltoid, to deltoid, often bidentate, leaf base attenuate extending into a broad petiole wing; petiole distinct, 23.6–31.0 × 3.1–3.3 mm, decurrent, channelled, often with a broadly sheathing base; upper stem leaves 50.0–68.0 × 4.4–12.2 mm, decreasing in size toward inflorescences, lamina narrowly lanceolate to linear-lanceolate, margin ± deeply and ± evenly incised, teeth in 6–14 widely and evenly spaced ± pairs, 1.5–7.2 mm long, protruding beyond leaf line, narrowly deltoid, tapering, acerose, ± acicular to acicular-falcate; lamina apex acute (rarely truncate-praemorse), acuminate, acumen 5.8–11.0 mm long, margins of acumen toothed, teeth often bidentate, 8–10 mm long, acerose, often acicular, or acicular-falcate, leaf base attenuate extending into a narrow petiole wing; petiole distinct, 2.9–8.2 × 1.2–3.3 mm, decurrent, channelled, often with a broadly sheathing base. Inflorescences racemose, 50–100 mm long at fruiting; rachis 0.5–2.25 mm diam., terminal and lateral, leaf-opposed, often long-persistent, sparsely to densely covered in pale, patent, ± clavate hairs or rarely glabrous, hairs 0.1–0.14 mm long; pedicels 5.6–7.2 mm long at fruiting, erecto-patent, with sparse, pale, patent, clavate hairs on adaxial surface, hairs 0.1–0.12 mm long. Flowers 3.0–4.5 mm diam., fragrant. Sepals 4, saccate, ± overlapping at base, lateral sepals broad, 0.5–1.5 mm diam., orbicular, pale to dark green with a broad white, ± undulose margin, apex rounded to obtuse, abaxial surface often hairy, hairs 0.2–0.4 mm long, eglandular or with glandular tip, mostly shedding at anthesis except near base, median sepals 0.5–0.9 mm diam., broadly elliptic, pale to dark green with a broad white, ± undulose margin, apex rounded to obtuse, abaxial surface glabrous. Petals white, 1.1–2.0 × 1.0–1.6 mm, spreading, claw 0.4–0.8 mm long; limb obovate, obovate-spathulate to orbicular, apex obtuse to rounded often slightly emarginate, margins smooth, sometimes weakly undulose. Stamens 4, filaments 1.2–2.0 mm long, white; anthers 0.3–0.5 mm long, yellow. Ovary 1.1–1.8 × 0.6–1.3 mm, ovate, broadly ovate to elliptic, green-brown, apex subacute; style 0.11–0.4 mm long, cylindrical; stigma 0.2–0.5 mm diam. Nectaries 4, 0.2–0.3 × 0.1–0.15 mm, narrow-oblong to deltoid, pale translucent green. Silicles cartilaginous when fresh, coriaceous when dry, 2.4–3.6 × 1.8–2.5 mm, elliptic to rhomboid, apex acute and tapering, valves green maturing grey-white, glabrous, scarcely separating at apex at maturity, not winged; style 0.3–0.7 mm long, exserted. Seeds 2, 1.3–1.9 × 0.8–1.6 mm, narrowly to broadly ovoid, brown to orange-brown, not winged. FL Jul–Jun. FR Sep–Jul.

Kermadec Islands: Curtis I., 18 July 1969, W. R. Sykes 836/K, (CHR 193789); Haszard Islet, 21 July 2002, T. Greene s.n., P. Dilks & P. Scofield, (AK 258039); Cheeseman Island, 19 November 1971, W. R. Sykes 932/K, (CHR 211838); Cheeseman Island, 26 July 2002, T. Greene s.n., P. Dilks & P. Scofield, (AK 258038);Cheeseman Island, “Te Mimi Paora Rock”, 24 May 2011, P. J. de Lange K850, (AK 326008); Cheeseman Island, “The Castle”, 24 May 2011, P. J. de Lange K851, (AK 326009); Cheeseman Island, “The Castle”, 24 May 2011, P. J. de Lange K852, (AK 326010). Cultivated (New Zealand): ex Curtis I. from CHR 193789, 9 January 1970, W. R. Sykes 901/K, (CHR 20188A); ex. Curtis I. from CHR 193789, 25 February 1971, W. R. Sykes 901/K, (CHR 201088B); Lincoln, ex Macauley Island, Landcare Research experimental nursery, 10 July 2008, P. B. Heenan s.n., (CHR 609796); Lincoln, ex Macauley Island, Landcare Research experimental nursery, 2 October 2009, P. B. Heenan s.n., (CHR 609806).

(Fig. 15). Endemic. Kermadec Islands (Macauley Island, Curtis Island, Haszard Islet and Cheeseman Island).

Lepidium castellanum is distinguished by its very robust, shrubby growth habit (Fig. 27), sometimes up to 1.8 × 2.0 m, erect, often closely packed, usually leafy stems, narrowly lanceolate to linear-lanceolate upper stem leaves, and by the very long, needle-shaped teeth of the leaves which reach well beyond the leaf margin (Figs 28–30). In this species, the pedicels (and often the inflorescence rachises) are minutely hairy (Fig. 31), whilst those of Lepidium oleraceum are glabrous. Lepidium castellanum is sympatric with Lepidium oleraceum on Curtis Island. Elsewhere, in northern New Zealand, especially on the Poor Knights and Hen (Taranga) islands, narrow-leaved forms of Lepidium oleraceum are also common. However, these plants have a much smaller growth habit, their leaves lack the distinctive long needle-shaped teeth and long-acuminate leaf apices of Lepidium castellanum, and, as with Lepidium oleraceum populations elsewhere, their pedicels are glabrous.

Lepidium castellanum is a sparse associate of the vegetation that has colonised the soft, erosion-prone cliff tops of Macauley Island, the less geothermally active summit slopes of Curtis Island, and the rocky tops of Haszard Islet and Cheeseman Island. On Macauley Island, Lepidium castellanum was recorded from open or sparsely vegetated, petrel-burrowed ground where it formed windswept shrubs in association with the creeping Scaevola gracilis Hook.f., tangles of Ipomoea cairica (L.) Sweet., Disphyma australe subsp. stricticaule Chinnock and Polycarpon tetraphyllum (L.) L. (Barkla et al. 2008; J. W. Barkla and T. C. Greene pers. comm.). On Curtis Island, it has been recorded growing amongst Disphyma australe subsp. stricticaule, while on the extensively seabird-burrowed Haszard Islet, it is one of the main shrub species, growing there in association with Myoporum rapense subsp. kermadecense (Sykes) Chinnock, Cyperus insularis Heenan et de Lange, Tetragonia tetragonioides (Pall.) Kuntze, and Parietaria debilis G.Forst. On Cheeseman Island, numerous seedlings and a few adults were seen in 2011 growing on heavily petrel-burrowed soil covered ledges amongst Disphyma australe subsp. stricticaule, and also as a few sub-adults within an active petrel colony in a narrow gully, growing in association with Cyperus insularis, Tetragonia tetragonioides, and Parietaria debilis.

Lepidium castellanum is confined to the Southern Kermadec Island group (see de Lange et al. 2004), part of the Kermadec Islands Nature Reserve. Within the southern Kermadec Islands group it has been recorded from Macauley Island, Cheeseman Island, Curtis Island, and from Haszard Islet. On Macauley Island it was recorded twice, in 2002 and 2006 (Barkla et al. (2008); T. C. Greene pers. comm.), at that time numbering at best five or fewer individuals. However, it was not seen on Macauley Island in 2011 (de Lange 2012; de Lange in press). In 2002 and 2006, the summit of Haszard Islet (6 ha, of which c.2.1 ha is vegetated) was thoroughly explored during helicopter-assisted landings by ornithologists. During those visits, it was noted that Lepidium castellanum was one of the major shrub-forming species on the richly manured and petrel-turbated soils of that islet’s summit slopes (T. C. Greene and K. Baird pers. comm.). While accurate counts of the number of plants present on Haszard Islet were not made, it would seem that on both visits the islet summit supported approximately 20–50 individuals (T. C. Greene and K. Baird pers. comm.). Further south, its status on Curtis Island remains unknown, as there have been no gatherings of Lepidium from there since 1969, and several visits during the 1980s and in 2002 did not see it there (G.A. Taylor and T. C. Greene pers. comm.). However, on nearby Cheeseman Island, a 7-ha island (of which c.5.6 ha is vegetated) two plants were recorded in 2002 during a brief helicopter assisted 40 minute visit (T. C. Greene pers. comm.). Later, in 2011, nine adults and an estimated 1000 seedlings were noted on Cheeseman Island during a thorough botanical investigation of that island (de Lange 2012). Therefore, to the best of our knowledge there are between 20 and 50 adults on Hazard Islet and nine mature plants on Cheeseman Island, while the status of this species on Curtis Island and Macauley Island is uncertain. Therefore Lepidium castellanum qualifies as ‘Threatened/Nationally Critical’ either because the total number of plants in the wild is < 250 (criterion A(1) of Townsend et al. (2008)) or because the combined area of occupancy is < 1 ha (criterion A(3) of Townsend et al. 2008). To that assessment we add the qualifiers ‘DP’ (Data Poor) and ‘IE’ (Island Endemic). Data Poor (‘DP’) is recommended because of the lack of accurate counts of wild plants from Haszard Islet, absence of any recent survey for the species on Curtis Island, and lack of any trend data.

A L. oleraceo habitu temporali, caulibus ad caudicem lignosum emorientibus, foliis crasse coriaceis, distincte petiolatis, late ellipticis, ellipticis vel obovatis, apicibus obtusis vel truncatis, marginibus saepe duplicato-serratis, siliculis orbiculatis vel orbiculato-rhomboideis, et sequentia nucleotidorum DNA distinguenda.

New Zealand (Fig. 33): Otago Land District, Otago Peninsula, Aramoana, Mole, open sites among stones and rocks at the edge of the road down mole and in car park, 26 November 2009, P. B. Heenan s.n., CHR 609777A! Isotype: CHR 609777B!

The specific epithet ‘crassum’ from the Latin for ‘thick’ refers to the distinctly thick and fleshy leaves of this species.

(Figs 34–37). Tap-rooted, strongly pungent smelling, perennial herb. Growth habit dense, stems closely placed, up to 50 cm tall, arising from underground woody stems. Stems upright to spreading, stout, short, rigid; mature stems woody, 100–400 × 10–12 mm, often devoid of foliage on middle and lower parts of stems, new stems 50–200 × 4–5 mm, leafy, glabrous. Leaves glabrous, coriaceous, green, undulate, rosette and stem leaves usually withering, variable in size and shape. Leaves of young and vigorous plants and stems: lamina 50–90 × 17–35 mm, broadly elliptic, elliptic to obovate; apex obtuse to truncate, often with up to 3 or 4 teeth; margin singly or doubly crenate, with 15–32 pairs of teeth; teeth up to 3.5 mm deep, sometimes overlapping, often protruding beyond leaf outline; base cuneate, petiole usually distinct; petiole up to 35.0 × 3.0–6.0 mm, channelled. Leaves of mature plants and cauline stems: lamina 15–45 × 6–15 mm, broadly elliptic, elliptic-obovate to obovate-oblong; apex obtuse to truncate, often with up to 3 or 4 teeth; margin singly crenate in upper half, teeth often protruding from leaf outline, with 5–10 pairs of teeth; teeth up to 1.2 mm deep, not overlapping, often protruding beyond leaf outline; base cuneate, sometimes narrowly so, usually tapering to ± distinct petiole; petiole 5–12 × 1.6–2.3 mm, channelled. Inflorescence terminal and lateral, racemose, 15–60 mm long, rachis 0.7–1.2 mm diam., glabrous; pedicels 4–7 mm long, erecto-patent, glabrous. Flowers 4.0–5.0 mm diam. Sepals 4, 1.3–1.6 mm long, saccate, overlapping at base, green, apex obtuse, margin white, shape dimorphic; lateral sepals broad, 1.4–1.5 mm diam., orbicular, abaxial surface often hairy, hairs 0.2–0.5 mm long; median sepals narrow, 1.0–1.3 mm diam., broadly elliptic, glabrous. Petals white, 1.8–2.0 × 1.0–1.1 mm, spreading, claw 0.6–0.8 mm long; limb broadly elliptic to orbicular, apex obtuse to rounded. Stamens 4; filaments 1.2–1.6 mm long, base 0.4–0.5 mm diam., equal; anthers 0.4–0.6 mm long. Ovary 1.0–1.6 × 0.9–1.1 mm, broadly ovate to broadly elliptic, green to green-brown, apex round or sometimes weakly shouldered; style 0.15–0.3 mm long, cylindrical; stigma 0.2–0.4 mm diam. Nectaries 4, 0.2–0.3 × 0.1–0.15 mm, oblong to oblong-triangular, green. Silicles cartilaginous when fresh, coriaceous when dry, 3.0–3.7 × 2.6–3.1 mm, orbicular to orbicular-rhomboid, apex obtuse to shallowly notched, valves pale brown, glabrous, not winged; style 0.2–0.3 mm long, exserted. Seeds 1.6–1.7 × 0.9–1.1 mm, narrowly ovoid, brown to orange-brown, not winged. FL Dec–Mar. FR Jan–Jul.

New Zealand (South Island): Oamaru, n.d., D. Petrie s.n., (WELT SP027632); Weston, North Otago, n.d., D. Petrie, s.n., (CHR 328313); south of Orore Point, North Otago, March 1982, T. R. Partridge s.n., (CHR 464024); Bridge Point, near Kakanui, North Otago, 13 February 1988, P. N. Johnson 751, (CHR 439956); Dunedin, February 1920, A. Wall s.n., (CHR 329217); Dunedin, St Leonards, 31 December 1896, B. C. Aston s.n., (WELT SP027641, SP027642); near Port Chalmers, Deborah Bay, March 1891, D. Petrie s.n., (WELT SP027633); Aramoana, The Mole, 13 November 1997, P. J. de Lange 3378 & J. W. Barkla, (AK 234312); Sandymount, Otago Peninsula, 12 May 1985, P. N. Johnson 300, (CHR 417850); Tairoa Head, Otago Peninsula, 27 March 1998, J. Barkla s.n. & L. Perriman, (CHR 579981); near mouth of Catlins River, n.d., D. Petrie s.n., (WELT SP027639); The Nuggets, South Otago, 5 Apr 1984, J. Ward s.n., (CHR 415751). South Otago, Waikawa Harbour, North Head, 9 February 1896, B. C. Aston s.n., (WELT SP027640); The Mole, Aramoana, 13 July 2009, P. B. Heenan s.n., (CHR 609772); Bridge Point, Orore Point, Waianakaru Road, 12 July 2009, P. B.Heenan s.n., CHR 609774. Cultivated (New Zealand): Auckland Regional Botanic Gardens, ex. Dunedin, The Mole, 1 December 1996, J. Lord 5/96, (AK 231116); Landcare Research experimental nursery, Lincoln, ex Bridge Point, 10 July 2008, P. B. Heenan s.n., (CHR 609797); Landcare Research experimental nursery, Lincoln, ex Aramoana, 10 July 2008, P. B. Heenan s.n., (CHR 609800); Landcare Research experimental nursery, Lincoln, ex Wharekakahu Island, 10 July 2008, P. B. Heenan s.n., (CHR 609801); Landcare Research experimental nursery, Lincoln, ex Nugget Point, 30 March 2009, P. B. Heenan s.n., (CHR 609805); Landcare Research experimental nursery, Lincoln, ex Wharekakahu Island, 25 January 2010, P. B. Heenan s.n., (CHR 609814).

(Fig. 15). Endemic. New Zealand, South Island, Otago and Southland. Lepidium crassum once ranged from the Waitaki Valley (an inland location) and coastal locations at Oamaru to North Head, Waikawa Harbour in the south Catlins. Today, Lepidium crassum is most common on Otago Peninsula, but occurs in small populations from near Kakanui, North Otago to The Nuggets, South Otago.

Lepidium crassum differs from the related Lepidium oleraceum by its usually much smaller stature (Fig. 34) and seasonal growth habit (with plants dying back to a basal rosette overwinter). Lepidium crassum has distinctly petiolate, uniformly broadly elliptic, elliptic to obovate, thickly coriaceous, often doubly crenate leaves with obtuse to truncate apices (Figs 35, 36). Its silicles are usually orbicular, sometimes orbicular-rhomboid, and with obtuse to shallowly notched apices (Fig. 37) and by its rDNA ETS sequence.

Known from scattered populations that usually comprise fewer than 50 plants in a small geographic area. Lepidium crassum usually occurs on coastal headlands and rocky outcrops where it grows in disturbed open areas and among coastal herbfield of Disphyma australe subsp. australe and Samolus repens (J.R.Forst. et G.Forst.) Pers. var. repens. This species has also colonised the man made ‘Mole’ at Aramoana (Fig. 34) where it is can be common amongst rubble, concrete blocks and landfill.

Lepidium crassum is known from a small number of wild populations on the Otago coast. As most populations comprise few plants or occur on unstable coastal headlands and rocky outcrops, this species is vulnerable to stochastic events. We estimate there are between 250 and 1000 plants and the largest population comprises fewer than 300 plants. Using the New Zealand Threat Classification System (Townsend et al. 2008), Lepidium crassum qualifies as ‘Threatened/Nationally Endangered’ (criterion B(1/1). We recommend appending the qualifiers ‘CD’ (Conservation Dependent – because there have been several populations established by the Department of Conservation and these are being actively managed), ‘DP’ (Data Poor – to reflect uncertainty over plant numbers), ‘EF’ (Extreme Fluctuations – because the largest known monitored population, that on Wharekakahu Island, appears to naturally undergo huge and apparently natural population fluctuations (J. W. Barkla pers. comm.)), and ‘RR’ (Range Restricted – because it is naturally confined to a geographically small part of the New Zealand Botanical Region).

New Zealand (North Island) (fide Garnock-Jones and Norton 1995): Onehunga, n.d. T.Kirk 341 WELT SP030080! Isolectotypes: WELT SP030089!, WELT SP027620!

The exact meaning of the species ‘flexicaule’ was not given by Kirk (1899) but the translation ‘flexuous stems’ accurately characterises the plant.

= Lepidium incisum Banks et Sol. ex Hook.f., Flora Novae-Zelandiae 1, 15 (1853) nom. illeg., non Roth, Neue Beytr. Bot. 1: 224 (1802), nec Edgew., Trans. Linn. Soc. 20, 33 (1851)

New Zealand (North Island) (fide Garnock-Jones and Norton 1995): Opuraga, on Beach, rare, n.d., Banks and Solander, BM! Isolectotypes: AK 100095!, WELT SP063696a, b!

Banks and Solander evidently chose the epithet ‘incisum’ in allusion to the deeply incised leaves of their specimens.

(Figs 38–43). Tap-rooted, strongly pungent smelling, decumbent, summer-green, perennial herb forming densely leafy, patches up to 1.5 m diam., and arising from stout, semi-circular, whitish-grey (when exposed) rootstock 3–10 mm diam. when fresh. Tap root fleshy, yellow to yellow-white when fresh, up to 300 mm long, deeply descending. Plants dying down in winter or in times of adversity to rootstock (in well-grown individuals the new season’s branchlets may die back to the previous season’s stem nodes). Stems prostate, weakly flexuous, divergent to widely spreading, 100–800 mm long, 0.8–5.2 mm diam., woody near base, initially spherical in cross-section, pale yellow-green to dark green, sometimes tinged maroon and finely puberulent with short papillate or tapered hairs; becoming glabrous and prominently ridged and/or grooved with age, and usually bearing numerous leaf scars and withered petioles; upper portion of stems sparingly and openly to heavily branched; branches and branchlets, usually very leafy. Leaves glabrous, firmly fleshy to succulent, glossy dark green to yellow-green, at senescence turning yellow. Rosette and lower stem leaves withering at fruiting; petioles distinct 10–50 × 1–3 mm, slightly concave in cross-section, fleshy; leaves glabrous above, with papillae or denticles along midrib beneath and margins in New Zealand plants, or without denticles (absent in Chatham Islands and most Australian plants); lamina 50–90 × 15–30 mm, pinnatifid, obovate to oblanceolate; pinnae in 1–3 pairs, bluntly toothed or crenate at apex and distal margins. Upper stem leaves glabrous above, usually with triangular denticles, dense on margins and sparse to dense beneath (although absent in Chatham Islands and most Australian plants); mostly apetiolate, petiole if present minute up to 1.5 mm long; lamina 5–30 × 3–10 mm, obovate, oblanceolate, or spathulate, apex bluntly toothed to crenate, base broadly to narrowly cuneate. Racemes 10–50 mm long, elongating up to 60 mm at fruiting, terminal and leaf-opposed; rachis and pedicels puberulent with short tapering hairs, and ± covered in triangular denticles (absent in Chatham Islands and most Australian plants) or glabrous; axillary; rachis and pedicels hairs if present, retrorse to patent, very short, 0.05–0.8 mm long, ± clavate, eglandular – glandular; pedicels, erecto-patent to patent, initially 1.04–1.27(–2.38) mm; elongating to 2.34–5.00(–6.02) mm long at fruiting. Flower buds dark green, apex bearing a conspicuous, caducous, crest of white, eglandular, antrorse hairs up to 0.9 mm long. Flowers sweetly fragrant, 1.4–1.8(–2.3) mm diam. Sepals, broadly ovate to oval, apex broadly obtuse, centrally green with a white margin, deeply concave, adaxially weakly keeled, adaxial midrib invested in conspicuous, caducous, white, eglandular, antrorse, hispid hairs, hairs sometimes scattered across rest of adaxial surface; abaxial surface glabrous; lateral sepals broad, 0.6–1.0 × 0.6–1.2 mm , median sepals narrower 0.4–0.8 × 0.4–0.7 mm. Petals white, 0.3–0.8(–1.0) × 0.2–0.8 mm, erecto-patent or patent, clawed; limb broadly obovate, apex obtuse, retuse or distinctly emarginate. Stamens 2, equal. Anthers c.0.16 mm long. Pollen bright yellow. Nectaries 4, subulate, 0.40 mm long. Silicles cartilaginous when fresh, subcoriaceous when dry, (2.1–)3.3(–4.1) × (2.2–)3.3–3.4(–4.0) mm, orbicular, obovate, to ovate, slightly winged in upper ¼, apex scarcely or shallowly notched, valves green maturing yellow-green, glabrous; style 0.8 (–1.0) mm long, free from the narrow wing, equal to or slightly exceeding the notch; stigma 0.2–0.4 mm diam. Seeds 2, 1.20–1.38 × 0.80–1.10 mm, ovoid to suborbicular, red-brown, dark red-brown or brownish black, not winged. FL. Oct–Feb. FR. Dec–Apr.

Australia (Tasmania): Port Davey, Gull Reef, January 1977, M. Allan s.n., (HO 26460); Port Davey, Elliot Point, 20 January 1986, A. Moscal 11876, (HO 402077); 1 km South of Rheuben Creek, 22 February 1985, A. Moscal 9734, (HO 401030); 1 km South of Nye Bay, 6 February 1986, A. M. Buchanan 8210, (HO 98828); Endeavour Bay, 30 January 1984, A. M. Moscal 6017, (HO 74481); Hibbs Bay, 25 January 1984, A. M. Buchanan 2775, (HO 74480); Bruny Island, n.d. L. Rodway s.n., (HO 54049). New Zealand (North Island): Great Barrier (Aotea island), Saddle Island, 7 January 1989, E. K. Cameron 5295, (AK 206897, AD, CHR).North Auckland, Waitemata, North Head, n.d., T. Kirk 342, (WELT SP030081); Waitemata, T. Kirk 41, n.d., WELT SP030088 (Duplicate AK 11429); Waitakere, May 1885, Ball s.n., (AK 261800); Bethell’s [Beach], 7 Jan 1934, L. M. Cranwell s.n., (AK 100102); Hauraki Gulf, Rangitoto Island, December 1882, T. F. Cheeseman s.n., (AK 4481); South Auckland, Manukau Harbour, Mangere, n.d., T. F.Cheeseman s.n., (AK 4480); Piako, n.d. J. Adams s.n., (AK 14865, AK 14866); Taranaki, Stent Road, 26 Jan 2010, P. J. de Lange 9279 & G. M. Crowcroft, (AK 317033), Kapiti Island, n.d. J. Buchanan s.n., (WELT SP087437). New Zealand (South Island): Between Little Whanganui River and Mokihinui, n.d., ?W. L. Townson 24, (WELT SP044035); Vicinity of Westport, 1903, W. Townson s.n., (WELT SP030082, WELT SP030084); Westport, Orowaiti River Bank, n.d., W. L. Townson s.n., (AK 253094); Westport, Orowaiti River Bank, 31 January 1953, R. Mason & N. T. Moar 2163, (AK 225200, CHR 81618); Cape Foulwind, n.d., W. Townson s.n., (WELT SP30085); Cape Foulwind, 4 February 1913, D. Petrie s.n., (WELT SP30086); Cape Foulwind, Tauranga Bay (Seal Colony), 14 August 1992, P. J. de Lange 1478 & D. A. Norton, (WELT SP079914); Punakaiki, 18 January 1931, L. B. Moore s.n. & L. M. Cranwell, (AK 100103). Chatham Islands: Rekohu, Point Somes, 25 January 2005, I. Keenan s.n., (AK 289897); Rekohu, Zimmermans Property, Point Somes, 10 January 2006, P. J. de Lange CH392, (AK 294940, CANB, CHR); Rekohu, Zimmermans Property, Point Somes, 10 January 2006, P. J. de Lange CH395, (AK 294940, CANB, CHR, WAIK); Rekohu, Ocean Bay, Unnamed Point south west of Bay, 14 January 2006, P. J. de Lange CH440 & J. W. D. Sawyer, (AK 295121); Rekohu, Wharekauri Farm Station, Cape Young, 13 January 2006, P. J. de Lange CH425 & J. W. D. Sawyer, (AK 295154).

(Fig. 44). Indigenous. Australia (Tasmania) and New Zealand (North, South, Chatham islands). In the North Island of New Zealand, Lepidium flexicaule is now known from only two extant populations, one on Saddle Island off the west coast of Great Barrier Island (Aotea Island) (de Lange and Cameron 2011) and the other at Stent Road north of Cape Egmont, Taranaki (Peet et al. 2003). Historically Lepidium flexicaule was present on Rangitoto Island, along the Waitakere Coastline, near North Head, Waitemata Harbour and near Onehunga, Manukau Harbour (Kirk 1899; Cheeseman 1906, 1925; Garnock-Jones and Norton 1995). It was also recorded from the Firth of Thames, at Mercury Bay (Kirk 1899; Cheeseman 1906, 1925; Garnock-Jones and Norton 1995) and near Wellington (Garnock-Jones and Norton 1995). In the South Island, the species is confined to the western coast of North-West Nelson from near Kaihoka to about Point Elizabeth. Lepidium flexicaule was discovered on the northern portion of Rekohu in 2005 by Department of Conservation botanists.

Lepidium flexicaule is easily distinguished from all other indigenous Lepidium species by the combination of having a decumbent growth habit, pinnatifid rosette and basal stem leaves (Figs 38–41), and flowers with two stamens. Most North Island and South Island populations are further distinguished by the presence of triangular denticles on branchlet stems, leaf margins and petioles, and on the rachis and pedicels of the inflorescences (Fig. 42, see also Garnock-Jones 1988). Denticles are, however, absent from the Chatham Islands, most Australian collections, and some South Island ones. The absence of denticles prompted the brief listing of Rekohu and Australian plants as an unnamed subspecies (see de Lange et al. 2009). However, as denticles are sometimes absent from South Island plants, formal taxonomic distinction may not be warranted. Nevertheless, it is worth noting that Rekohu plants are consistently larger than those seen from the rest of this species’ range, with well-grown specimens reaching up to 1.5 m diam., and that cpDNA data separates Tasmanian and Rekohu Lepidium flexicaule from New Zealand samples (Fig. 4). Further study, including obtaining sequences of non-denticulate South Island Lepidium flexicaule, is warranted. Of the indigenous species, it is perhaps most similar to Lepidium naufragorum, which differs by its upright bushy shrub-forming habit with ascending to erect stems (Fig. 55), more sharply serrated pinnae of the leaves; racemes that are not so clearly leaf-opposed and are much longer (up to 150 mm, cf. 50 mm long), flowers that have four rather than two stamens, and emarginate petals that are longer than the sepals (see Garnock-Jones and Norton (1995) for other distinctions).

Lepidium flexicaule is however, commonly confused in the field with the naturalised Lepidium didymum L. and Lepidium coronopus (L.) Al-Shehbaz, species with which it sometimes grows. Both Lepidium didymum and Lepidium coronopus have long been treated in New Zealand under the segregate genus Coronopus but Al-Shehbaz et al. (2002) reinstated both species within Lepidium. Lepidium flexicaule can be distinguished from Lepidium didymum and Lepidium coronopus by the valves of the silicles, which are smooth and which dehisce into 2 valves to leave a persistent replum. The silicles of Lepidium didymum and Lepidium coronopus have a reticulate-ridged or warty surface, and do not dehisce along the valves; in Lepidium didymum they split into 1-seeded segments and in Lepidium coronopus they are indehiscent.

The ecology of Lepidium flexicaule was described in some detail by Garnock-Jones and Norton (1995). They concluded that it is a strictly coastal species of turf communities, rock crevices and the strandline of bouldery beaches. Lepidium flexicaule has also colonised tracksides, the compacted peaty ground of seal colonies, bird roosts, and even bird nests, where plants presumably arose from fruiting material that had been used for nest construction. In most locations, the species is found within the spray zone and always in sites prone to frequent disturbance.

Previously Lepidium flexicaule had been assessed ‘Threatened / Nationally Vulnerable _{CD, EF}’ (de Lange et al. 2009). Based on current evidence, this ranking is no longer appropriate, as Rekohu populations of this species (treated in that paper as an unnamed entity Lepidium aff. flexicaule but now included in Lepidium flexicaule (see recognition above)) and all of the monitored New Zealand Lepidium flexicaule populations are in decline, while the past total area of occupancy had been grossly under estimated. Based on current knowledge, Lepidium flexicaule is more appropriately assessed as ‘Threatened / Nationally Endangered’ (criterion A(3/1) of Townsend et al. (2008)) because the total area of occupancy of the sum of the populations is < 10 ha, and evidence obtained from monitoring indicates that there is an ongoing decline predicted to be up to 50% over the next ten years due to increased plant mortality (causes of which are as yet unknown) and loss of habitat caused by competition from weeds and coastal erosion. To this assessment we append the qualifiers ‘CD’ (Conservation Dependent – because virtually all known populations are being managed), ‘EF’ (Extreme Fluctuation – because monitoring also indicates that this is a species that, despite the decline, is naturally prone to seasonal population fluctuations) and ‘TO’ (Threatened Overseas – because, for the time being, we include Rekohu and Tasmanian plants within Lepidium flexicaule (see recognition), and in Tasmania Lepidium flexicaule is also seriously threatened (A. Buchanan pers. comm.)).

On the Chatham Islands, Lepidium flexicaule is occasionally found growing with Lepidium oligodontum and Lepidium rekohuense, and putative hybrids have been collected when these species occur together. There is good evidence for the hybrid Lepidium flexicaule × Lepidium oligodontum (represented by AK 294939, P. J. de Lange CH391; AK 294942, P. J. de Lange CH394; AK 295119, P. J. de Lange CH442 & J. W. D. Sawyer). This hybrid is known from two sites on Chatham (Rekohu) Island where it is found in association with both parents. Like the parents, the hybrids have a decumbent growth habit. They share with Lepidium flexicaule pinnatifid rosette leaves, but vegetative features of Lepidium oligodontum are evident in the mid to upper stem leaves that are mostly spathulate to cuneiform (but with occasional deeply lobed margins to weakly pinnatifid leaves that are similar to Lepidium flexicaule). The flowers of the hybrids have variable stamen numbers ranging from 1–5 per flower, compared to 2 in Lepidium flexicaule and 2–4–6 in Lepidium oligodontum, while pollen stainability varied from 80 to 85%. Seedlings raised from AK 295119 presented a bewildering array of foliage types grading into either parent. Unfortunately, through mishap these plants failed to reach maturity, so vouchers showing this are unavailable.

Evidence for Lepidium flexicaule × Lepidium rekohuense is less convincing. This putative hybrid is known from only one gathering (AK 295155, P. J. de Lange CH426 & J. W. D. Sawyer) which was collected in January 2006 from Cape Young, Rekohu, at a site where both parents grew. In the field, this specimen resembled Lepidium flexicaule closely except that the plant was distinctly leafy and the upper stem foliage was more copious and larger than is usual for Lepidium flexicaule. As with all Rekohu gatherings of Lepidium flexicaule, this specimen lacked stem and marginal leaf denticles. Furthermore, as with both parents, there are two stamens per flower, and pollen stainability of this gathering was 98%. No fruiting material was present, and on a subsequent search of this site in 2007 the putative hybrid and its parents had gone. This specimen may be a very well grown example of Lepidium flexicaule but, as it appears anomalous alongside other Rekohu, New Zealand, and Australian gatherings of that species, we prefer to treat it as a putative Lepidium flexicaule × Lepidium rekohuense hybrid.

The putative hybrid Lepidium flexicaule × Lepidium oleraceum has also been recorded once, as a spontaneous plant arising in cultivation at the Auckland Regional Council Botanic Gardens (AK 228296 S. P. Benham s.n., AK 223492 S. P. Benham s.n.). The hybrid grew in a site where both parents (Scots Beach Lepidium flexicaule and Stephens Island Lepidium oleraceum) were planted together. The hybrid appeared from a seed lot gathered from the cultivated Lepidium flexicaule plant. It initially appeared to be different because it was much larger and more vigorous than Lepidium flexicaule, which it otherwise closely resembled (S. P. Benham pers. comm.). Over time, the hybrid developed weakly ascendant branches with rhomboid deeply serrated to pinnatifid mid and upper stem leaves, and flowers with 1–6 stamens. The hybrid appeared to be fully fertile, and seedlings raised from it showed clear segregation back to either parent (S. P. Benham pers. comm.). Again through mishap, these plants were lost before they reached maturity and specimens could be taken.

A L. oleraceo habitu decumbente vel effuso patuloque, caulibus sparsis effusis gracilibus flexilibus, foliis subcoriaceis, apice obtuso vel truncato, plerumque 2--3 dentibus prominentibus ut videtur irregularibus, floribus fructibusque e plantis juvenibus prodientibus, et propria sequentia nucleotidorum DNA distinguenda.

New Zealand (Fig. 45): Otago Land District, Otago Peninsula, Long Beach, Purakanui, on stabilised sand at base of cliff amongst introdued grasses, 26 November 2009, P. B. Heenan s.n., CHR 609785B! Isotype CHR 609785A!

The specific epithet ‘juvencum’ from the Latin for ‘youthful’ refers to plants flowering and fruiting only a few months after germinating from seed.

(Figs 46–49). Tap-rooted, strongly pungent smelling, perennial herb. Growth habit open, straggly, up to 50 cm tall. Stems usually decumbent to sprawling, slender, flexible, sparse; mature stems woody, 100–1000 × 8–12 mm, often devoid of foliage on middle and lower parts of stems; new stems 100–400 × 3–4 mm, leafy, glabrous. Leaves glabrous, subcoriaceous, green, often undulate, rosette and stem leaves usually withering, variable in size and shape. Leaves of young and vigorous plants and stems: lamina 37–87 × 12–32 mm, elliptic, obovate or elliptic-oblanceolate; apex truncate to obtuse, usually with 2–3 prominent teeth and often appearing irregular; margin singly crenate, with 4–19 pairs of teeth; teeth up to 1.5 mm deep, not overlapping; base attenuate to cuneate, tapering to a distinct or indistinct petiole; petiole up to 23.0 × 2.0–5.0 mm, or sessile. Leaves of mature plants and cauline stems: lamina 10–60 × 3–21 mm, elliptic, elliptic-oblanceolate, obovate to elliptic-obovate; apex subacute, truncate or obtuse, usually with 2–3 prominent teeth and often appearing irregular; margin singly crenate in upper and/or lower half, with 4–19 pairs of teeth; teeth up to 1.3 mm deep, not overlapping; base attenuate to cuneate, tapering to distinct or indistinct petiole, or sessile. Inflorescence terminal and lateral, racemose, 10–60 mm long, rachis 1.0–1.3 mm diam., glabrous; pedicels 4.0–6.0 mm long, erecto-patent, usually glabrous although lower pedicels occasionally sparsely hairy on adaxial surface. Flowers 4.0–5.0 mm diam. Sepals 4, 1.3–1.5 mm long, saccate, overlapping at base, green, apex obtuse, margin white, shape dimorphic; lateral sepals broad, 1.1–1.5 mm diam., orbicular, abaxial surface often hairy, hairs 0.1–0.5 mm long; median sepals narrow, 0.9–1.2 mm diam., broadly elliptic, glabrous. Petals white, 2.0–2.4 × 1.1–1.5 mm, spreading, claw 0.6–1.0 mm long; limb broadly elliptic to orbicular, apex obtuse to rounded. Stamens 4(–5); filaments 1.4–1.7 mm long, base 0.3–0.5 mm diam., equal; anthers 0.3–0.4 mm long. Ovary 1.0–1.5 × 0.9–1.4 mm, broadly ovate to broadly elliptic, green to green-brown, apex usually with shoulders; style 0.15–0.25 mm long, cylindrical; stigma 0.3–0.5 mm diam. Nectaries 4, 0.2–0.3 × c. 0.1 mm, oblong-obovate, green. Silicles cartilaginous when fresh, coriaceous when dry, 3.1–4.2 × 2.5–3.5 mm, elliptic-rhomboid to orbicular-rhomboid, valves light brown, glabrous, apex shallowly notched, not winged; style 0.2–0.3 mm long, exserted. Seeds 1.6–1.8 × 0.9–1.3 mm, narrowly ovoid, brown to orange-brown, not winged. FL Nov–May. FR Nov–Jun.

New Zealand (South Island): Otago, Long Beach, 30 December 1998, G. S. Stone s.n., (AK 239841); Otago, Long Beach, 25 Jun 1997, D. Nelson s.n. & J. Barkla, (CHR 518601); Otago, Long Beach, 13 July 2009, P. B. Heenan s.n., (CHR 609779);Otago, Long Beach, 14 July 2009, P. B. Heenan s.n., (CHR 609780); Tomahawk Beach, 4 February [1920?], B. C. Aston s.n., (WELT SP027651, WELT SP052362); Tavora Reserve, 13 July 2009, P. B. Heenan s.n., (CHR 609786–609789);Otago Peninsula, Green Island, January 1955, R. A.. Falla s.n., (WELT SP038744); Otago Peninsula, Green Island, January 1957, M. E. Gillham s.n., (CHR 111452); Otago Peninsula, Green Island, 6 July 1983, P. N. Johnson 70, (CHR 404351); Otago Peninsula, Green Island, 27 March 1989, P. N. Johnson 823, (AK 185642, CHR 461272); Otago Peninsula, Green Island, 12 February 1999, G. Loh s.n., (AK 238646); Solander Island, 10 November 1973, P. N. Johnson s.n., (CHR 253066). New Zealand (Stewart Island/Rakiura): Halfmoon Bay, Mill Bay, 4 February 1963, P. Hynes s.n., (AK 92212); Womans Island, January 1976, P. N. Johnson s.n., (CHR 261892); Herekopare Island, 20 May 1959, E. A. Willa s.n., (WELT SP050477, AK 249215); Herekopare Island, 4 January 2010, P. B. Heenan s.n, (CHR 609808–609810); Mutton Bird (Titi) Islands, Big Moggy Islands, December 2002, B. D. Rance s.n., (AK 293254). Cultivated (New Zealand): Lincoln, ex Herekopare Island, 2 Feb 1980, H. D. Wilson 798-314, (CHR 368843); Lincoln, ex Long Beach, Landcare Research experimental nursery, 10 July 2008, P. B. Heenan s.n., (CHR 609803).

(Fig. 44). Endemic. New Zealand. South and Stewart Island/Rakiura. Known from Otago and islands off the coast of Stewart Island/Rakiura Lepidium juvencum is currently known from just three locations: Long Beach and Green Island (Otago Peninsula) and Herekopare Island (east of Stewart Island/Rakiura). However, it is also probably present on some of the south-western Titi Island (south-west of Stewart Island/Rakiura) (B.D. Rance pers. comm.) and it is still likely to occur elsewhere, especially on rat free islands in the vicinity of Stewart Island, as is evidenced by specimens previously collected from Womans (CHR 261892) and Solander (CHR 253066) islands in the 1960s and 1970s.

Lepidium juvencum is distinguished by open, sprawling, straggling growth habit, and the stems are often trailing on the ground (Figs 46, 47). The leaves are elliptic, elliptic-oblanceolate, obovate to elliptic-obovate, and have attenuate bases; the marginal teeth are usually small and are mostly confined to the distal third of the leaf (Fig. 48). The silicles are most like Lepidium oleraceum but differ by possessing a rounded, slightly notched apex (Fig. 49).

Lepidium juvencum grows in open and often disturbed sites on sandy soil. At Long Beach it grows on stabilised sand at the base of a cliff immediately behind the beach, and at Herekopere Island it occurred on a steep coastal bank comprising sandy loam. On Kaimohu Island, in the south-western Titi group, what is probably Lepidium juvencum has been recorded growing with Lepidium limenophylax on the margins of coastal scrub. Lepidium juvencum has also been established and is self-seeding at the Yellow-eyed Penguin Trust revegetation project at Tavora Reserve (near Palmerston, North Otago). Here it grows among Euphorbia glauca G.Forst. and Ficinia spiralis (A.Rich.) Muasya et de Lange on sand dunes at the back of the beach.

Lepidium juvencum is presently known from three wild localities, and, while the number of plants at these localities is not known with certainty, it is estimated that there are < 100 in total. It is also acknowledged that Lepidium juvencum is likely to be more widespread, especially in the vicinity of Stewart Island/Rakiura, and further field surveys are required. Recruitment is likely to be sporadic, and the small and sparse populations are especially vulnerable to stochastic events, such as storm surges, that could erode its habitat. Based on our current knowledge of this species and using the New Zealand Threat Classification System (Townsend et al. 2008), Lepidium juvencum qualifies as ‘Threatened/Nationally Critical’ (either criterion A(1) or A(3) of Townsend et al. (2008) applies). We recommend appending the qualifiers, ‘CD’ (Conservation Dependent – because the species has been deliberately established at Tavora Reserve, where it is actively managed), ‘DP’ (Data Poor – to reflect uncertainty over its distribution), and ‘RR’ (Range Restricted – because it is naturally confined to a geographically small part of the New Zealand Botanical Region).

A L. oleraceo G.Forst. habitu constanter decumbenti vel procumbenti, nodis caulorum ligneis repullantibus tempo novo et interdum radicibus adventiis, foliis constanter angusto-lanceolatis vel linearibus serrulatis floribus staminibus duobus, et DNA sequentiis singularibus differt.

New Zealand (Fig. 50): Snares Island, Snares Island Expedition 1969/72, University of Canterbury, December 1969, C. H. Hay s.n., WELT SP077578!

The specific epithet ‘limenophylax’ derives from Greek (Ό λιμνοφυλαξ) and, though it strictly means ‘lake watcher’ (C. Blackford pers. comm.), it is used here in the sense of the alternative meaning of “harbour watcher or coast-guard”, which Green (1994; p. 535) states is the more appropriate derivation (see his comments on the etymology of the Norfolk Island endemic orchid Phreatia limenophylax (Endl.) Benth.). It is given in allusion to the preferred habitat of Lepidium limenophylax on the Snares, i.e. the low turf confined to the exposed ends of rocky headlands jutting out into and overlooking the sea. Historically, it was on these exposed headlands that the World War II coastwatchers were posted to keep watch for signs of enemy shipping (Fleming 1948, McEwen 2005).

(Figs 51–54). Tap-rooted, strongly pungent smelling, summer-green, perennial herb, arising from stout rootstock 7.3–10.0 mm diam. Plants dying down to rootstock and previous season’s stem nodes during winter. Stems decumbent, bases often very stout, ± spherical, somewhat woody when mature 10–15 × 10–12 mm, composed of numerous old leaf and stem bases, sometimes producing roots, new season’s grow decumbent, spreading 10–300 × 3–10 mm, glabrous, sometimes with very sparse, appressed, caducous, silky 0.5–1.0 mm long, hairs near stem apices, at fruiting, often devoid of foliage from much of length. Leaves glabrous, firmly fleshy to succulent, usually dark green to green, sometimes yellow-green. Rosette and stem leaves usually withering at fruiting but sometimes with a few long persistent. Petiole distinct, 20–40 × 2–4 mm lamina 50–100 × 5–15 mm, lanceolate, narrowly lanceolate to linear lanceolate; distal ⅓–⅔ finely but sharply serrated or crenate, teeth in 10–20 pairs running to and including subacute, obtuse to rounded apex (teeth not extending beyond leaf outline), base cuneate to narrowly cuneate. Middle stems leaves with petiole indistinct, lamina narrowly linear lanceolate to linear, often recurved to falcate in the distal ⅓ – ½ of leaf length, 50–120 × 3–6 mm; upper ⅔ or occasionally the entire leaf finely but sharply serrated; teeth in 12–30 pairs running to and including the apex, and not extending beyond leaf outline, lamina base tapered, very narrowly cuneate. Upper stem leaves with or without a distinct petiole, petiole if present 40–60 mm, linear to linear-spathulate, occasionally narrowly lanceolate, often recurved or falcate from ½ of leaf length, 30–100 × 2.0–30.0 mm, margins finely but sharply serrated. Racemes 5–15 mm long, terminal and axillary; rachis glabrous; pedicels glabrous, erecto-patent, 2–8 mm long at fruiting. Flowers c. 0.4–1.0 mm diam. Sepals 4, saccate, overlapping at base, green with pale-green to white thickened margin, apex broadly obtuse, shape and size dimorphic; lateral sepals c.0.6–1.0 × 0.6–1.2 mm, broadly ovate to oval, mostly glabrous, sometimes sparsely hairy, hairs 0.2–0.4 mm long, caducous; median sepals 0.9–1.2 × 0.5–1.2 mm, elliptic to obovate, abaxial surface sparsely hairy, hairs 0.2–0.4 mm long, caducous. Petals white to off-white, 1.5–2.0 × 0.3–1 mm, erecto-patent to somewhat spreading, clawed; limb narrowly obovate, apex obtuse, occasionally emarginated. Stamens 2, equal. Nectaries 2, subulate, 0.35 mm long. Silicles cartilaginous when fresh, coriaceous when dry, 2.5–3.5 × 1.5–3.3 mm, elliptic or rhomboid, not winged, apex usually notched (rarely truncate), valves green maturing yellow-green, glabrous; style 0.1–0.3 mm long, exceeding the shallow notch (if notch present); stigma 0.3–0.5 mm diam. Seeds 2, narrowly ovoid, brown, red-brown to orange-brown, not winged, 1.25–1.3 × 0.35–0.60 mm. FL. Nov–Feb. FR. Nov–Feb.

New Zealand (Stewart Island/Rakiura): Kaimohu Island, 25 February 1965, B. A. Fineran s.n., (CANU 8703). Snares Islands: November 1907, B.C. Aston s.n., (AK 4464, WELT SP027621, WELT SP027628); January 1909, B. C. Aston s.n., (WELT SP027623, WELT SP027624, WELT 27625, WELT SP027626); west coast between Signpost Hill and Razor Back, 29 November 1984, G. S. Hardy s.n., (WELT SP078810); on headland just below Signpost Hill, 29 November 1984, G. S. Hardy s.n., (WELT SP078811); December 1947, n. c., R. C. Murphy Expedition, (WELT SP035432); December 1947, F. Newcombe s.n., (WELT SP09710); cliff edge, February 1961, B. Fineran s.n., (CANU 5949); cliff edge, February 1961, B. A. Fineran s.n., (CANU 5950); B. Fineran s.n., Feb 1961, cliff edge, CANU 5994; cliff edge, February 1961, B. A. Fineran s.n., (CANU 5995); cliff edge, February 1961, B. A. Fineran s.n., (CANU 6000A–6000B); Broughton Island, 4. November 1972, D. S Horning s.n., (CANU 18916); July 2002, P. Sagars.n., (AK 283482). Auckland Islands: January 1890, T. Kirk s.n., (WELT SP027634).

(Fig. 44). Endemic. New Zealand, South-western Titi (Muttonbird) Islands (Kaimohu Island), The Snares (North East, Broughton islands), and Auckland Islands. Lepidium limenophylax is possibly also present on Pohowaitai Island (B. A. Fineran, 4 Mar 1965, CANU 8723) another of the south-western Titi (Muttonbird) Islands, although this collection, comprising sterile specimens of a whole young plant and a branch are inadequate to be certain. There are also additional unconfirmed observations of what may be this species from many of the other south-western Titi (Muttonbird) Islands (B.A. Fineran and B.D. Rance pers. comm.).

Lepidium limenophylax is recognised by the decumbent growth habit (Figs 51, 52), with plants developing a distinct woody network of branches. New season’s growth arises from the nodes left from the previous season’s growth. In this species, the leaves are consistently lanceolate, narrowly lanceolate, linear lanceolate or linear (Figs 50, 51B, 52), though slightly broader in seedlings and basal rosettes. The flowers have two stamens (Fig. 53). It is morphologically most similar to Lepidium rekohuense, which also has a decumbent growth habit and flowers with two stamens. Lepidium limenophylax has lanceolate, narrowly lanceolate, linear lanceolate or linear leaves, rather than the spathulate to narrowly ovate or elliptic leaves of Lepidium rekohuense. Furthermore, the silicles of Lepidium limenophylax are elliptic, rhomboidal and unwinged (Fig. 54), rather than orbicular, obovate to ovoid and winged.

On the Snares, Lepidium limenophylax has been described as an occasional component of cliff top vegetation where it is stated to grow in association with living or dead Poa astonii Petrie pedestals (and on occasion in association with Poa tennantiana Petrie and Hebe elliptica (G.Forst.) Pennell), often near the feeding sites of brown skuas (Stercorarius lonnbergi Mathews, 1912) and occasionally around the nests of Buller’s mollymawk (Thalassarche bulleri (Rothschild, 1893)) or on dry peaty cliff tops (sometimes within the sandy quartz derived from the underlying granite) (Fineran 1969, Hay et al. 2004, Lake and Evans 2011). Further inland, it was also reported by Fineran (1969, p. 3) ‘as uncommon... sometimes present on sites of old abandoned penguin rockeries, where it usually grew with great vigour’. Brian Rance (in litt.), on a visit to the islands in 2000, found that, as with Fineran (1969), Lepidium limenophylax was virtually confined to the north-western cliffs. More recently, Lake and Evans (2011) failed to find it at any inland sites. All these authors and individuals noted Lepidium limenophylax as uncommon in other habitats, though a few plants have been noted near Boat Harbour growing in association with Poa astonii, Asplenium obtusatum G.Forst.and Hebe elliptica in semi-open situations caused by Hooker’s sea lions (Phocarctos hookeri (Gray, 1844)) (Lake and Evans 2011; B. D. Rance pers. comm.). Very little is known about the associations of Lepidium limenophylax on Kaimohu Island, though herbarium notes suggest that it grew there in association with what is probably Lepidium juvencum (see comments under that species), and B. A. Fineran (pers. comm.) noted that this species was present mostly along ridge crests and above cliff faces, always in association with the feeding sites of skua. Brian Rance (in litt.) has also observed plants apparently matching Lepidium limenophylax on several of the Titi (Muttonbird) islands south west of Stewart Island, usually growing along tracks in association with a sparse ground cover of Apium prostratum subsp. prostratum var. filiforme (A.Rich.) Kirk, Nertera depressa Banks et Sol. ex Gaertn and Poa astonii under semi-open Olearia angustifolia Hook.f. / Olearia lyallii Hook.f. short forest.

On the Snares Lepidium limenophylax occupies a very narrow habitat range scattered over four key areas and covering an area of 943 m² (Lake and Evans 2011). Lake and Evans (2011) were unable to count or estimate population size stating that ‘it was not possible to estimate the number of plants due to the tendency [of the species] to grow in patches and not [as] isolated plants’. Nevertheless, they reported ‘a good mix of mature and juvenile plants’. Outside these islands, the status of this species on the Auckland and south-western Titi (Muttonbird) Islands is unknown. Previously, Lepidium limenophylax, as Lepidium aff. oleraceum (c) (CANU 5995; Snares), had been listed by de Lange et al. (2009, p. 89) under Appendix 2 Taxonomically Indeterminate Listings, as ‘Acutely Threatened/Nationally Critical’ with the qualifiers ‘IE’ (Island Endemic) and ‘RR’ (Range Restricted) appended. That assessment used data provided by B. D. Rance, on a visit to the Snares Islands in 2000. Lake and Evans (2011) provided a detailed update on the information obtained by Rance during a three week stay on the Snares between September and October 2010. They concluded that the species was in good health, and much more abundant than had been believed previously. While these observations suggest that this is probably a narrow-range species secure within its Snares stronghold, we recommend retention of the current threat listing of de Lange et al. (2009) as a precautionary measure because: 1. Accurate information about the status of this species outside the Snares is absent (i.e. it is not a Snares endemic), 2. The total area of occupancy on the Snares is < 1 ha (criterion A(3) in Townsend et al. (2008)) and 3. There is still no trend data available for the species. However, to reflect the observations of Lake and Evans (2011) and our ongoing uncertainty about population trend and the status of this species outside the Snares we recommend that the addition of the qualifier ‘DP’ (Data Poor) to the current threat listing. It should also be noted that because Lepidium limenophylax is known from a number of islands and island groups it does not meet the definition of ‘IE’ in Townsend et al. (2008). Lepidium limenophylax also should be qualified ‘CD’ (Conservation Dependent) because the largest known population occurs within a Nature Reserve and World Heritage site (The Snares) where it is vulnerable to the spread of disease, weed and rat incursions, all of which would have a profound impact on its long-term security. Because of these threats, which put at risk not only the Lepidium but all other Snares terrestrial biota, access to the islands is strictly controlled by the New Zealand Department of Conservation, which also undertakes regular biosecurity inspections of the island group.

New Zealand: Open Bay Islands, Taumaka, 15 February 1992, P. J. Garnock-Jones 2121, D. A. Norton & D. R. Given, CHR 470212!

The epithet ‘naufragorum’, the genitive plural form of ‘naufragus’ meaning a ‘castaway or ship wrecked person’, was given by Garnock-Jones and Norton (1995) to commemorate a sealing gang that was set down on the type locality of the species, the Open Bay Islands, in 1810 where they were left for four years until they were rescued.

(Figs 55, 56). Tap-rooted, pungent-smelling, summer-green, perennial herb forming a densely leafy, shrub up to 680 mm tall, rootstock stout, 3–10 mm diam. when fresh semi-circular, whitish-grey (when exposed). Tap root fleshy, yellow to yellow-white when fresh, up to 300 mm long, deeply descending. Plants dying down in winter or in times of adversity to rootstock. Stems ascending to erect, 150–680 mm long, glabrous, 2–8 mm diam., woody near base, prominently ridged and/or grooved with age, and usually bearing numerous leaf scars and withered petioles, pale yellow-green to dark green, sometimes tinged maroon; mid to upper portion of stems much branched; branches and branchlets, usually very leafy. Leaves glabrous, fleshy to ± subcoriaceous, glossy dark green to yellow-green, at senescence turning yellow. Rosette and lower stem leaves withering at fruiting; petioles distinct 40–80 × 1–3 mm, slightly concave in cross-section, fleshy, winged; lamina 60–140 × 15–25 mm, pinnatifid (rarely simple), narrow-oblong to narrow-oblanceolate; pinnae when present in 3–8 pairs, sharply toothed toothed at apex and distal margins, lamina of simple leaves, deeply and unevenly serrated, teeth 1.0–2.2(–5.1) mm long, usually not projecting beyond leaf outline. Middle stem leaves similar (rarely simple) or becoming shallowly pinnatifid, sharply serrate. Upper stem leaves 10–50 × 2–10 mm, narrow-obovate to linear oblanceolate, pinnatifid to simple, sharply toothed at apex and at apex of pinnae if present, cuneate at base; petiole minute or absent. Racemes 10–120 mm long, elongating up to 140 mm at fruiting, terminal and axillary, ± leaf-opposed; rachis glabrous or sparsely hairy; pedicels sparsely hairy, erecto-patent, initially1.3–1.6 mm; elongating to 3–5 mm long at fruiting. Flower buds grass green to dark green, apex glabrous or sometimes bearing a conspicuous, caducous, crest of white, eglandular, antrorse hairs up to 0.9 mm long. Flowers sweetly fragrant, 2.8–3.2 mm diam. Sepals, ovate to oval, apex broadly obtuse, green with a white scarious margin, deeply concave, adaxially weakly keeled, abaxial midrib glabrous or invested in conspicuous, caducous, white, eglandular, antrorse, flexuous hairs, hairs sometimes scattered across rest of abaxial surface; adaxial surface glabrous; lateral sepals 0.8–1.2 × 0.8–1.2 mm , median sepals narrower 0.7–1.0 × 0.6–0.9 mm . Petals white, 0.9–1.4 × 0.6–0.9 mm, erecto-patent, clawed; limb obovate, apex emarginate. Stamens 4, equal. Anthers c.0.12 mm long. Pollen bright yellow. Nectaries 4, subulate, 0.25 mm long. Silicles cartilaginous when fresh, subcoriaceous when dry 2.8–4.0 × 2.3– 3.2 mm, broadly elliptic, slightly winged, apex shallowly notched, valves green maturing pale green, glabrous; style 0.1–0.2 mm long, free from the narrow wing, equal to or slightly exceeding the notch; stigma 0.4 mm diam. Seeds 2, 1.6–2.0 × 1.2–1.8 mm, obovoid, orange-brown, not winged, mucilaginous when wet. FL. Oct–Mar. FR. Nov–Apr.

New Zealand (North Island): Auckland City, Mt Albert, Jesmond Terrace (Naturalised), 9 January 1998, P. J. de Lange 3415, (AK 234450). New Zealand (South Island): North Westland, Punakaiki, Seal Island, 29 December 1991, D. A. Norton s.n., (CANU 36438); Punakaiki, Perpendicular Point, 13 February 1992, P. J. Garnock-Jones 2112 & D. A. Norton, (CHR 470203); Motukikie, January 1995, G. Loh s.n., (CANU 37018); Paringa Knight’s Point, 1 January 1990, D. A. Norton s.n. & J. M. Lord, (CANU 35508); Open Bay Island, January 1909, B. C. Aston s.n., (WELT SP030114); Open Bay Islands, Taumaka Island, 15 February 1992, P. J. Garnock-Jones 2117, (AK 229863, CHR 470208); Open Bay Islands, Taumaka Island, 15 February 1992, P. J. Garnock-Jones 2118, D. A. Norton & D. R. Given, (CHR 470209); Open Bay Islands, Taumaka Island, March 2006, P. I. Knightbridge s.n., (AK 317068—pinnate-leaved form); Open Bay Islands, Taumaka Island, 4 January 2006, P. I. Knightbridge s.n., (AK 317070—simple-leaved form).

(Fig. 44). Endemic. New Zealand, South Island, (Westland) where it is known from seven sites from Cape Foulwind south to the Open Bay Islands (Taumaka and Popotai). With the exception of the Open Bay Islands, Lepidium naufragorum is scarce within the mainland part of its range. Lepidium naufragorum has also been collected once as a gutter weed in Auckland City, plants having established there from a nearby garden where the species was being cultivated. These naturalised plants persisted for about five years before they died out.

Lepidium naufragorum is easily distinguished from all other Lepidium species in New Zealand by the upright bushy shrub habit (with plants dying down to a central root stock in winter), erect stems, mostly sharply serrated pinnatifid rosette and stem leaves (rarely entire) (Fig. 55), and by the emarginate silicles (Fig. 56). Of the New Zealand species, it is most similar to Lepidium flexicaule in which it was included by Garnock-Jones (1988) before field work and further study recognised its distinctiveness. For differences between Lepidium naufragorum and Lepidium flexicaule, see under Lepidium flexicaule.

Although most plants of Lepidium naufragorum have pinnatifid foliage, occasional specimens with simple or weakly pinnatifid leaves are also known, and these can at times be locally common (Knightbridge and Newton 2006). It was plants such as these that were the basis for literature records (e.g. Burrows 1972) and collectors such as L. Cockayne and B.C. Aston (WELT!) referring this species to Lepidium oleraceum and Lepidium banksii (as var. ovatum). From Lepidium oleraceum these forms can easily be distinguished by the seasonal growth habit (with plants dying back to the rootstock over winter), and by the distinctly emarginate silicles. So far, Lepidium oleraceum has not been found within the Westland range of Lepidium naufragorum. The confusion with Lepidium banksii partly stems from these early collectors’ uncertainty as to what this species is. In any case, non-pinnate leaved forms of Lepidium naufragorum are readily distinguished from Lepidium banksii by their much smaller (2.8–4.0 × 2.3– 3.2 mm) broadly elliptic and shallowly notched silicles. The silicles of Lepidium banksii are much larger (4.5–5.5 × 4.0–5.0 mm), broadly ovate and deeply notched (Fig. 25).

The ecology of Lepidium naufragorum was described in some detail by Garnock-Jones and Norton (1995) who noted that it preferred sites frequented by seals and nesting sea birds or sea bird roosts. They concluded that the species required these species not only for nutrient enrichment but also because their disturbance kept the habitats of this species open. The authors also noted that Lepidium naufragorum was restricted to base-rich substrates.

Lepidium naufragorum has a conservation status of “Threatened / Nationally Vulnerable _{CD, RR}” (de Lange et al. 2009). Based on current evidence this ranking is still appropriate.

A L. panniformo habitu effuso, foliis saepe nitentibus sparse leviter dentatis ab apicibus surculorum confertis et a serie rDNA ETS differt. A. L. oleraceo habitu effuso aestati-virenti, siliculis valde incisuris et a serie rDNA ETS differt.

Chatham Islands (Fig. 57): Chatham Is., Mangere Island, Mangere Island Nature Reserve, South of Hut Landing, 14 February 2006, P.J. de Lange CH876 & P.B. Heenan, AK 300342! Isotypes: CHR 552303A-C!

The epithet “oblitum” from the Latin meaning “forgotten, disregarded or neglected” refers to the circumstances in which this species was first discovered. At an early stage of this revision Lepidium oblitum had been included within the range of Lepidium panniforme, with some plants treated as possible hybrids between Lepidium panniforme and Lepidium oleraceum. It was only following critical analysis of plants grown in New Zealand from samples collected from Mangere Island that the distinctive nature of this species was realized using DNA and morphological data.

(Figs 58–62). Tap-rooted, strongly pungent, sprawling, laxly though much-branched, leafy perennial shrub, forming patches up to 1.0 × 0.9 m. Tap-root up to 700 mm long, ± napiform, branched 1–2 or more times. Rootstock 6–14 mm diam., woody, exposed portion ± smooth, often covered in dried leaf remnants and withered branch stubs. Plants usually dying down to rootstock and/or previous seasons stem nodes, towards end of growing season. Stems mostly seasonal, a few ± persistent (i.e. lasting 2 or more years), arising from rootstock base and basal portion of main central stem, widely and unevenly spaced or closely packed, woody, lax and sprawling, sometimes arching and subascendent, weakly angled to ± terete, glabrous; mature stems 2.1–4.6 mm diam., 0.3–0.8 m long; green to yellow-green, brittle, bases bearing numerous leaf abscission scars, otherwise devoid of leaves for at least the first ⅔ with the final ⅓ distinctly leafy and often much branched when vegetative, with the leaves tending to fall at flowering and fruiting; upper stems similar, though distinctly leafy and pliant. Leaves coriaceous, fleshy, green to dark green, often glossy. Rosette leaves 5–14(–20), mostly present in autumn – early spring usually not persisting (very rarely so) at fruiting; petioles distinct, up to 40 × 2 mm, flat or slightly concave in cross-section, succulent; lamina oblanceolate, cuneiform, obovate-oblong to spathulate up to 60 × 22 mm, margins finely to deeply incised in upper ¼–⅓, teeth in 3–6(–8) pairs running to praemorse apex, basal few teeth pair usually asymmetric, base narrowly attenuate. Middle stem leaves mostly persisting at fruiting; petiole up to 32 mm long, mostly flat in cross-section, sometimes slightly concave, usually winged; lamina narrowly oblanceolate, oblanceolate, spathulate, oblong, obdeltoid, 15.0–32.0(–56.2) × 12–16.8(–18.6) mm; margins finely to deeply incised in distal ¼–⅓, teeth in 3–6(–8) pairs running to the usually praemorse apex, basal few teeth pairs usually asymmetric, lamina base narrowly attenuate, extending as a wing (0.8–1.2 mm wide) usually to petiole base. Upper stem leaves petiolate, petiole 12 mm long, flat or slightly concave, usually broadly winged; lamina narrowly oblanceolate, narrowly lanceolate, narrowly spathulate, to linear-cuneiform, 10.2–20.0(–29.0) × 2.4–3.5(–4.8) mm; margins entire or deeply dentate, if dentate then usually asymmetrically tridentate (rarely with 1–4 teeth), lamina base narrowly attenuate, extending as a wing (0.3–1.7 mm wide) almost to petiole base. Racemes (30–)50(–76) mm long, usually congested, elongating up to 100 mm at fruiting, terminal and axillary; rachis and pedicels glabrous (pedicels very rarely bearing a few minute, caducous, glandular hairs near base); pedicels, erecto-patent to patent, 0.4–0.8(–1.0) mm long, elongating to 0.8–1.2(–2.0) mm long at fruiting. Flower buds dark green to green, apex glabrous. Flowers sweetly fragrant, 1.6–1.8(–2.0) mm diam. Sepals 4, saccate, pale to dark green with a broad white, ± undulose margin, pale to dark green with a broad white, ± undulose margin, deeply concave, adaxially weakly keeled or not; lateral sepals 0.9 × 0.6 mm, broadly ovate to oblong, ± overlapping at base, apex rounded to obtuse, adaxial surface glabrous (sometimes diffusely papillate), abaxial surface usually glabrous, sometimes hairy near base, hairs patent, weakly flexuous, 0.1–0.3 mm long, eglandular, shedding at anthesis; median sepals 1.0 × 0.9 mm, broadly ovate to oblong, apex rounded to obtuse, adaxial surface glabrous, abaxial surface glabrous. Petals overtopping sepals, white, 0.9–1.15(–1.3) × 0.5–0.6(–0.8) mm, patent, clawed; limb broadly obovate, apex weakly retuse. Stamens 2–4, equal. Anthers c. 0.10 mm long. Pollen bright yellow. Nectaries 2, subulate, 0.3–0.42 mm long. Silicles cartilaginous when fresh, coriaceous when dry, orbicular, orbicular-ovate to ± rhomboid, (2.8–)3.0(–3.3) × (2.4–)2.6(–3.0), margin winged, notably more so toward apex, apex notched, base obtuse, valves green maturing yellow-green, glabrous, dried surface ± coarsely reticulate; style 0.1–0.2(–0.3) mm long, free from the narrow wing, usually exceeding the notch; stigma 0.18–0.22 mm diam, capitate. Seeds 2, ovoid to ellipsoid, orange-brown to dark red-brown, not winged, 1.8–1.9 × 0.9–0.93 mm. FL Nov–Mar. FR Jan–Jun.

Chatham Islands: Rabbit Island, 14 February 2006, P.J. de Lange CH878 & P.B. Heenan, (AK 300344). Mangere Island, Mangere Island Nature Reserve, Above Hut, 14 February 2006, P.J. de Lange CH874 & P.B. Heenan, (AK 300340; CHR 552373); Mangere Island, Mangere Island Nature Reserve, ‘Top Plateau’, 22 February 2011, B. Gibb s.n., (AK 331585).

(Fig. 63). Endemic. New Zealand, Chatham Is., Mangere and Rabbit Islands.

Lepidium oblitum is recognised by its sprawling growth habit (Fig. 58); by the branches often widely spaced, trailing and mostly devoid of foliage except for the ends which are typically densely terminated by foliage (Fig. 57); and by the prominently notched silicles (Fig. 62). Of those species with which it grows (Lepidium oligodontum, Lepidium oleraceum, Lepidium panniforme and Lepidium rekohuense), it can be easily distinguished from Lepidium oleraceum by its sprawling growth habit, flowers with 2–4 stamens, and distinctly notched silicles (Figs 58, 61, 62).

From Lepidium oligodontum, Lepidium oblitum is easily distinguished by it’s more upright, though somewhat, untidy, sprawling growth habit (Fig. 58), and longer lived, persistently leafy branches. The foliage of Lepidium oblitum (Figs 59, 60) has leaf margins that are consistently toothed (never sparingly-toothed or entire), while the flowering racemes are distinctly longer ((30–)50(–76) mm cf. (5.0)–9.7(–28.9) mm in Lepidium oligodontum)) and on fruiting they may elongate up to 100 m (rather than 60 mm). In Lepidium oblitum the flowers have 2–4 rather than 2–4–6 stamens (Fig. 61B, 61C), while the silicles are never fleshy or turgid, rather they are distinctly cartilaginous and prominently (rather than weakly) notched (Fig. 62). Lepidium oblitum also has consistently shorter styles ((0.1–0.2(–0.3) mm long) than Lepidium oligodontum ((0.2–0.8(1.2) mm long)).

Lepidium oblitum and Lepidium panniforme grow together on Mangere Island (Fig. 63). As they have similar silicles and flowers with 2–4 stamens but very different foliage, Lepidium oblitum was initially thought to be hybrid between Lepidium panniforme and Lepidium oleraceum. At that time it was not realised how widespread Lepidium oblitum is, nor that it comprised stable, true-breeding populations (G. Houliston unpubl. data). It was only following DNA-based investigations undertaken for this paper that the putative hybrid status was rejected and its status as a species recognised. Nevertheless, Lepidium oblitum is still closely allied to Lepidium panniforme, and both also share a relationship to the extinct Lepidium obtusatum, a relationship evident by all three species sharing prominently notched silicles. Lepidium oblitum has widely spreading, lax (rather than erect to suberect), sprawling stems and rosette leaves that distinguish it from Lepidium panniforme (Figs 82, 83). The leaves of Lepidium oblitum are shallowly toothed but they are never as deeply serrated or lacerate as those of Lepidium panniforme (Figs 81, 83, 84). DNA data suggests that gene flow between Lepidium oblitum and Lepidium panniforme has occurred at some sites on Mangere Island (G. Houliston unpubl. data).

Lepidium oblitum also grows with Lepidium rekohuense on Rabbit Island. From that species it is easily separated by its smaller growth habit; by the glabrous rather than sparsely papillate-hairy upper branch stems, and glabrous rather than finely hairy inflorescences (the pedicels of Lepidium oblitum may, very rarely, bear a few caducous glandular hairs). Furthermore, the flowers of Lepidium oblitum have 2–4 rather than 2 stamens, while the silicles of Lepidium rekohuense are also mostly orbicular (rarely obovate) and slightly larger (up to 4.1 × 4.0 mm cf. 3.3 × 3.0 mm).

Lepidium oblitum is one of three Lepidium species recorded from Mangere and Rabbit islands. On these islands, it is frequently found growing in association with petrel burrows, and on exposed, often wind eroded cliff faces, associated soil blowouts and ephemeral drainages and seepages. On Mangere Island, it is perhaps most common among the boulders and rocks forming the northern summit of the Top Plateau. In these habitats, Lepidium oblitum commonly grows with Asplenium obtusatum, Festuca coxii (Petrie) Hack., Disphyma papillatum Chinnock, Puccinellia chathamica (Cheeseman) Allan et Jansen and Senecio radiolatus F.Muell. subsp. radiolatus. On Mangere Island, it is also associated with Aciphylla dieffenbachii (F.Muell.) Kirk, Hebe chathamica (Buchanan) Cockayne et Allan, Hebe dieffenbachii (Benth.) Cockayne et Allan (and hybrids between these two Hebe), Myosotidium hortensium (Decne.) Baill., and, on occasion, Lepidium oleraceum and Lepidium panniforme. On Rabbit Island it is associated with Atriplex australasica Moq., Critesion murinum (L.) Á.Löve subsp. murinum, Malva arborea L., Lepidium oligodontum and Lepidium rekohuense.

Lepidium oblitum is so far known only from the Chatham Islands group where it has been collected from Mangere and Rabbit Islands (Fig. 63). On Mangere Island (a Nature Reserve, with strict permit controlled access) it is known from several populations, the largest of which occurs along the northern western cliff faces of ‘Top Plateau’. That population may number in the tens of plants; the only other ones known occur in steep runnels draining the western cliffs south of the main Landing area. When visited by PdL and PBH in 2006, there were fewer than 10 adults in total. Subsequent visits to Mangere Island by Department of Conservation rangers suggest that there has been little change in the population sizes on that island. However, hard data is unavailable and so it would inappropriate to infer from these observations that Lepidium oblitum populations on Mangere Island are stable. On nearby privately owned Rabbit Island, two plants of Lepidium oblitum were seen in 2006 and the island has not been visited by Department of Conservation staff since.